IRLF 


^a 


LI  BR  ARY 

OF   THE 

UNIVERSITY  OF  CALIFORNIA. 

GIF'T    OF" 


M 

Received. ...TOT .28   1892        ,  /^  81OLOGY 

,  LIBR 

?.  LfC(QO  h      Shelf  No.  . 


—3 
fc 


THE 


MAMMOTH    CAYE 


AND  ITS 


INHABITANTS, 

OR  DESCRIPTIONS  OF   THE 

FISHES,   INSECTS   AND    CRUSTACEANS 

FOUND   IN   THE   CAVE  ; 


WITH      FIGURES      OF      THE     VARIOUS      SPECIES,     AND     AN    ACCOUNT     OF 
ALLIED    FORMS,  COMPRISING   NOTES    UPON   THEIR   STRUC- 
TURE,   DEVELOPMENT    AND    HABITS,    WITH 
REMARKS    UPON    SUBTERRANEAN 
LIFE    IN    GENERAL. 


A.  S.  PACKARD,  JR.,  AND  F.  W.  PUTNAM, 


EDITORS   OF  THE  AMERICAN  NATURALIST. 


SALEM  : 
NATURALISTS'  AGENCY. 

1872. 


OP  THB 

tf  HIV  EH  SIT  7 


00 


PRINTED  AT  THE 

SALEM     PRESS. 

Corner  of  Liberty  and  Derby  Streets, 

SALEM,  MASS. 
F.    W.   PUTNAM    &   CO. 


PREFACE. 


THE  following  pages  were  first  published  in  the  "  Ameri- 
can Naturalist"  for  December,  1871  and  January,  1872,  with 
the  exception  of  the  Synopsis  of  the  family  including  the 
Blind  fishes  of  the  cave,  which  was  first  published  in  the 
"Annual  Keport  of  the  Peabody  Academy  of  Science  for 
1871." 

In  bringing  the  several  articles  together  in  the  present 
form  but  slight  changes  have  been  made,  principally  in  the 
form  of  a  few  additional  notes. 

It  will  undoubtedly  be  the  good  fortune  of  some  visitors 
to  the  cave  to  discover  other  kinds  of  animals  than  those 
mentioned  in  the  following  pages,  and  to  observe  new  facts 
relating  to  the  habits  of  the  various  species.  For  it  must 
be  remembered  that  all  the  observations  thus  far  recorded 
have  been  made  by  but  a  very  few  of  the  thousands  who 
annually  visit  the  Mammoth  Cave,  and  that  no  thorough  zo- 
ological exploration  of  the  cave  has  yet  been  undertaken. 
Should  any  new  facts  be  observed,  or  unknown  species  dis- 
covered, the  authors  of  this  little  work  would  be  pleased  to 
be  informed  of  them,  and  communications  on  all  such 
matters  are  solicited  for  publication  in  the  pages  of  the 

AMERICAN  NATURALIST. 

THE  AUTHORS. 

PEABODY  ACADEMY  OF  SCIENCE,  Salem,  Mass.,  Feb.  1872. 


TJHI7WSITY 


THE  MAMMOTH  CAVE 


AND  ITS 


INHABITANTS. 


CHAPTER   I. 

THE  FORMATION   OF   THE   CAVE.* 

BY  F.    W.    PUTNAM. 

AFTER  the  adjournment  of  the  meeting  of  the  American  Associ- 
ation for  the  Advancement  of  Science,  held  at  Indianapolis,  in 
August  last,  a  large  number  of  the  members  availed  themselves  of 
the  generous  invitation  of  the  Louisville  and  Nashville  Railroad 
Company,  to  visit  this  world  renowned  cave,  and  examine  its  pe- 
culiar formation  and  singular  fauna. 

The  cave  is  in  a  hill  of  the  subcarboniferous  limestone  forma- 
tion in  Edmondson  County,  a  little  to  the  west  and  south  of  the 
centre  of  Kentucky.  Green  river,  which  rises  to  the  eastward  in 
about  the  centre  of  the  state,  flows  westward  passing  in  close 
proximity  to  the  cave,  and  receiving  its  waters  thence  flows  north- 
westerly to  the  Ohio. 

The  limestone  formation  in  which  the  cave  exists,  is  a  most  in- 
teresting and  important  geological  formation,  corresponding  to 
the  mountain  limestone  of  the  European  geologists,  and  of  con- 
siderable geological  importance  in  the  determination  of  the  west- 
ern coal  fields. 

We  quote  the  following  account  of  this  formation  from  Major  S. 
S.  Lyon's  report  in  the  fourth  volume  of  the  Kentucky  Geological 
Survey,  pages  509-10. 

"The  sinks  and  basins  at  the  head  of  Sinking  creek  exhibit 
in  a  striking  manner,  the  eroding  effect  of  rains  and  frost  —  some 
of  the  sinks,  which  are  from  forty  to  one  hundred  and  ninety  feet 

*From  the  AMERICAN  NATURALIST  for  December,  1871. 

(5) 


FORMATION    OF    THE    CAVE. 

deep,  covering  an  area  of  from  five  acres  to  two  thousand.  The 
rim  of  sandstone  surrounding  these  depressions  is,  generally, 
nearly  level ;  the  outcropping  rocks  within  are  also  nearly  horizon- 
tal. Near  the  centre  there  is  an  opening  of  from  three  to  fifteen 
feet  in  diameter  ;  into  this  opening  the  water  which  has  fallen  within 
the  margin  of  the  basin  has  been  drained  since  the  day  when  the 
rocks  exposed  within  were  raised  above  the  drainage  of  the  coun- 
try, and  thus,  by  the  slow  process  of  washing  and  weathering,  the 
rocks,  which  once  filled  these  cavities,  have  been  worn  and  carried 
down  into  the  subterranean  drainage  of  the  country.  All  this  has 
evidently  come  to  pass  in  the  most  quiet  and  regular  manner. 
The  size  of  the  central  opening  is  too  small  to  admit  extraordinary 
floods  ;  nor  is  it  possible,  with  the  level  margin  around,  to  suppose 
that  these  cavities  were  worn  by  eddies  in  a  current  that  swept  the 
whole  cavernous  member  of  the  subcarboniferous  limestone  of 
western  Kentucky  ;  but  the  opinion  is  probable  that  the  upheaving 
force  which  raised  these  beds  to  their  present  level,  at  the  same 
time  ruptured  and  cracked  the  beds  in  certain  lines ;  that  after- 
wards the  rains  were  swallowed  into  openings  on  these  fractures, 
producing,  by  denudation,  the  basins  of  the  sinkhole  country,  and 
further  enlarging  the  original  fractures  by  flowing  through  them, 
and  thus  forming  a  vast  system  of  caverns,  which  surrounds  the 
western  coal  field.  The  Mammoth  Cave  is,  at  present,  the  best 
known,  and,  therefore,  the  most  remarkable." 

So  much  has  been  written  on  the  cave  and  its  wonders,  that  to 
give  a  description  of  its  interior  would  be  superfluous  in  this 
connection,  even  could  we  do  so  without  unintentionally  giving 
too  exaggerated  statements  which  seems  to  be  the  natural  result 
of  a  day  underground,  at  least  so  far  as  this  cave  is  concerned, 
for  after  reading  any  account  of  the  cave,  one  is  disappointed  at 
finding  the  reality  so  unlike  the  picture.  As  the  Association  party 
was  accompanied  by  one*  who,  while  a  most  enthusiastic  collector 
and  explorer,  was  also  a  calm  recorder  of  statements  made  by 
the  geologists  of  the  party,  we  cannot  do  better  in  conveying  to 
our  readers  the  general  geological  character  and  structure  of  the 
cave  than  to  copy  his  account. 

"  As  we  expected  to  remain  within  the  cave  a  long  time,  our 
trusty  guide,  Frank,  had  provided  himself  with  a  well-filled  can 
of  oil,  to  replenish  our  lamps,  and  with  this  strapped  upon  his 
back  he  led  the  way  into  the  thick  darkness.  We  shall  attempt 
no  description  of  the  cave.  Its  darkness  must  be  felt  to  be  ap- 
preciated, and  no  form  of  expression,  understood  by  mortals  who 
have  never  descended  to  its  cavernous  depths,  nor  trod  its  gloomy 

*  W.  P.  FJSHBA.CK,  Esq.,  of  the  Indianapolis  Daily  Journal. 


FORMATION    OF    THE    CAVE.  / 

corridors,  can  convey  anything  like  an  adequate  idea  of  the  place. 
After  spending  fifteen  hours  within  its  chambers,  it  is  absolutely 
nauseating  to  read  the  descriptions  which  have  been  current  in  the 
letters  of  newspaper  correspondents  for  a  quarter  of  a  century, 
and  even  the  vigorous  and  picturesque  language  of  Bayard  Taylor 
becomes  tame  and  commonplace  when  it  attempts  to  describe  this 
subterranean  wonder  of  the  world. 

How  and  when  the  cave  was  made,  were  the  leading  questions 
in  the  minds  of  the  geologists.  They  do  not  believe  that  the  cave 
was  the  immediate  result  of  some  violent  upheaval  of  the  strata, 
which  left  these  vast  crevices  and  chambers  of  which  the  cave  is 
composed  ;  neither  do  they  share  the  popular  belief  that  the  rapid 
and  violent  action  of  some  subterranean  stream  of  water  has 
worn  these  deep  channels  through  the  limestone  ;  on  the  contrary, 
they  find  conclusive  evidence  that  the  same  agencies  are  at  work 
and  the  same  changes  in  progress  to-day  that  have  been  slowly, 
steadily  and  quietly,  through  vast  periods  of  time,  accomplishing 
the  marvellous  wonders  that  now  astonish  the  beholder.  The  cave 
is  wrought  in  the  stratum  known  as  the  St.  Louis  limestone,  which 
in  some  places  reaches  a  thickness  or  depth  of  four  hundred  feet. 
This  stone  is  dissolved  whenever  it  is  subjected  to  the  influence  of 
running  or  dripping  water  impregnated  with  carbonic  acid  gas. 
Water  exposed  to  the  air  readily  absorbs  this  gas,  and  surface  water 
percolating  through  small  fissures  of  the  limestone,  dissolves  it. 
Another  fact  should  be  stated.  When,  during  this  process  of  so- 
lution, the  water  becomes  thoroughly  impregnated  with  lime,  it 
loses  its  power  to  dissolve  the  stone.  The  following  conditions, 
then,  were  essential  to  the  productions  of  the  cave,  assuming  what 
is  not  disputed  by  geologists,  that  the  place  where  the  cave  now  is, 
was  once  nearly  solid  limestone.  First,  that  there  should  be  fis- 
sures in  the  strata,  allowing  the  ingress  of  the  surface  water.  Sec- 
ondly, there  should  be  a  place  or  places  of  exit  for  the  water  charged 
with  limestone  in  solution.  Without  the  latter,  the  water  would 
become  charged  with  lime,  fill  up  the  crevices,  and  the  dissolving 
process  would  cease.  These  conditions  are  all  present  to-day,  and 
have  remained  the  same  during  the  countless  ages  that  have  passed 
away  while  the  work  has  been  in  progress.  There  have  doubtless 
been  times  in  the  history  of  the  cave,  when,  owing  to  a  greater 
flow  of  water,  the  work  has  progressed  more  rapidly  than  at  pres- 
ent, but  that  the  results  have  been  accomplished  in  the  manner 
stated,  rather  than  by  the  process  of  attrition  by  rapid  currents  of 
large  volumes  of  water,  seems  to  be  the  general  opinion  of  scien- 
tific men.  This  theory  is  strengthened  by  the  fact  that  where  the 
cave  attains  its  greatest  heights,  and  reaches  its  lowest  depths, 
the  dripping  waters  have  never  ceased  their  labors,  and  are  busily 
at  work  to-day.  In  the  Mammoth  Dome,  for  instance  —  rarely 
seen  by  visitors,  on  account  of  the  dangers  and  fatigue  incident  to 
the  journey  —  where  the  chasm  attains  a  height  and  depth  of  more 


8  FORMATION    OF    THE    CAVE. 

than  two  hundred  and  fifty  feet,  a  cascade  falls  from  a  great  height, 
and  keeps  the  entire  surface  of  the  rocks  covered  with  dripping 
water.  This,  falling  into  a  deep  pit  below,  finds  an  exit  through 
which  it  bears  away  a  portion  of  the  lime  composing  the  rock. 
After  a  walk  of  thirteen  hours,  our  guide  informed  us  that  he  would 
conduct  us  to  the-  Mammoth  Dome  if  we  felt  able  to  bear  the 
fatigue  of  the  journey.  Foot-sore  and  weary,  we  were  not  in  a 
favorable  condition  for  so  arduous  an  undertaking,  but  Mr.  Thomas 
Kite  of  Cincinnati,  who  had  visited  the  locality  thirty  years  ago, 
urged  us  to  go,  and  told  us  the  sight  of  this  Dome  was  worth  all 
the  rest.  Provided  with  magnesium  and  calcium  lights,  we  crawled 
and  climbed  our  way  to  the  brink  of  the  pit,  the  bottom  of  which 
was  reached  by  a  rickety  ladder,  slippery  and  dripping  with  water. 
A  portion  of  the  party  descended,  and  when  all  were  ready  the 
lights  were  ignited,  and  the  immense  dome  was  revealed  to  us  in 
all  its  majestic  beauty.  Upon  our  return,  three  hearty  cheers  were 
given  to  the  good  friend  at  whose  earnest  solicitation  we  under- 
took this  part  of  our  journey. 

We  are  indebted  to  Professor  Alexander  Winchell.  of  the  Unir 
versity  of  Michigan,  for  the  following  abstract  of  his  views  con- 
cerning the  formation  of  the  cave. 

'  The  country  of  the  Mammoth  Cave  was  probably  dry  land  at 
the  close  of  the  coal  period,  and  has  remained  such,  with  certain 
exceptions,  through  the  Mesozoic  and  Csenozoic  ages,  and  to  the 
present.  In  Mesozoic  times,  fissures  existed  in  the  formation, 
and  surface  waters  found  their  way  through  them,  dissolving  the 
limestone  and  continually  enlarging  the  spaces.  A  cave  of  con- 
siderable dimensions  probably  existed  during  the  prevalence  of  the 
continental  glaciers  over  the  northern  hemisphere.  On  the  dis- 
solution of  the  glaciers,  the  flood  of  water  which  swept  over  the 
entire  country,  transporting  the  materials  which  constituted  the 
modified  drift,  swept  through  the  passages  of  the  cave,  enlarging 
them,  and  leaving  deposited  in  the  cave,  some  of  the  same  quart- 
zose  pebbles  which  characterize  the  surface  deposits  from  Lake  Su- 
perior to  the  Gulf  of  Mexico.  Since  the  subsidence  of  the  waters 
of  the  Champlain  epoch,  the  cave  has  probably  undergone  compar- 
atively few  changes.  The  well  one  hundred  and  ninety-eight  feet 
deep,  at  the  further  end  of  the  cave,  shows  where  a  considerable 
volume  of  the  excavatory  waters  found  exit.  The  Mammoth  Dome 
indicates  probably,  both  a  place  of  exit  and  a  place  of  entrance 
from  above.  So  of  the  vertical  passages  in  various  other  portions 
of  the  cave.' 

We  believe  that  the  views  of  Professor  Winchell  are  in  harmony 
with  those  of  the  other  eminent  geologists  of  the  party,  and  when 
it  is  considered  that  the  geologists  of  this  excursion  stand  in  the 
front  rank  of  the  most  eminent  scientific  men  of  the  world,  their 
views  upon  this  interesting  subject  are  well  Avorthy  of  attention. 
Before  dismissing  this  branch  of  the  subject,  we  will  take  occasion 


FORMATION    OF   THE    CAVE. 

to  correct  a  popular  error  concerning  the  formation  of  the  beauti- 
ful structures  that  adorn  the  ceilings  of  some  portions  of  the  cave. 
In  the  dryer  localities,  where  the  floors  are  dusty  and  everything 
indicates  the  prolonged  absence  of  moisture,  the  ceiling  is  covered 
with  a  white  efflorescence  that  displays  itself  in  all  manner  of 
beautiful  shapes.  It  requires  no  stretch  of  the  imagination  to  dis- 
cover among  these,  the  perfect  forms  of  many  flowers.  The  lily 
form  prevails,  and  the  ceilings  of  many  of  the  chambers  are  cov- 
ered with  this  beautiful  stucco  work,  surpassing  in  delicacy  and 
purity  the  most  beautiful  workmanship  of  man.  These  are  not 
produced,  as  many  suppose,  by  the  dripping  of  water,  and  the 
gradual  deposit  of  sulphate  of  lime  upon  the  outer  portions.  The 
stalactite  is  formed  in  this  manner,  but  these  are  neither  stalacti- 
form,  nor  are  they  produced  in  a  similar  way.  Dripping  water 
is  the  agency  that  forms  the  stalactite,  while  the  efflorescence 
in  the  dryer  portions  of  the  cave  cannot  take  place  where  there  is 
much  moisture.  The  growth  of  these  beautiful  forms  is  from 
within,  and  the  outer  extremities  are  produced  first.  They  are 
the  result  of  a  sweating  process  in  the  limestone  that  forces  the 
delicate  filaments  of  which  they  are  composed  through  the  pores 
upon  the  surface  of  the  rock,  their  beautiful  curved  forms  result- 
ing from  unequal  pressure  at  the  base,  or  friction  in  the  apertures 
through  which  they  are  forced.  Mr.  L.  S.  Burbank,  of  Lowell, 
Mass.,  has  kindly  furnished  us  with  the  following  abstract  of  his 
opinions  upon  this  interesting  subject. 

'  The  rosettes,  wreaths,  and  other  curved  fibrous  forms  of  gyp- 
sum, in  the  Mammoth  Cave,  occur  only  in  particular  strata  of  the 
limestone  which  do  not  appear  in  the  first  part  of  the  long  route. 

Their  formation  may  be  explained  in  this  way :  that  portion  of 
the  rock  where  they  are  found  consists  of  carbonate  of  lime,  with 
some  impurities,  and  contained  originally  the  sulphide  of  iron,  or 
iron  pyrites,  disseminated  in  small  grains  or  crystals,  and  also  in 
rounded  nodules  or  concretions,  sometimes  of  considerable  size. 

By  exposure  to  air  and  moisture,  oxygen  unites  with  both  the 
sulphur  and  the  iron,  producing  sulphuric  acid  and  oxide  of  iron, 
which  combined,  form  a  sulphate  of  iron.  Then  a  double  de- 
composition takes  place ;  the  sulphuric  acid  unites  with  the  lime 
to  form  the  gypsum  ;  the  carbonic  acid  of  the  limestone  combines 
with  the  oxide  of  iron,  forming  a  carbonate  of  iron,  and  this,  on 
further  exposure,  parts  with  the  carbonic  acid,  and  leaves  the 
brown  coating  of  oxide,  which  is  seen  in  many  places  on  the  sur- 
face of  the  rock. 

The  gypsum  is  thus  constantly  forming  in  the  rock,  and,  being 
soluble,  is  carried  by  the  water  to  the  exposed  surface  where  it 
crystallizes. 

The  crystals  appear  to  grow  out  from  the  rock  by  additions  from 
beneath,  which  continue  to  push  the  ends  first  formed,  and  if  these 
do  not  become  attached  to  other  parts  of  the  rock,  straight  needle- 


10  ANIMAL    LIFE    IN    THE    CAVE. 

like  fibres  are  often  produced.  Very  commonly,  however,  the 
crystals  begin  to  form  when  a  small  nodule  of  the  iron  ore  is  ex- 
posed at  the  surface ;  the  parts  first  formed  become  attached  to 
the  surface  around  the  edges,  and  as  the  chemical  action  proceeds 
towards  the  centre  of  the  nodules  successive  leaf-like  layers  are 
thrown  out,  and  the  rosette  form  is  the  result.  Along  lines  of 
fracture  in  the  surface  of  the  rock,  the  crystals  are  curved  in  op- 
posite directions. 

The  wreaths  and  other  figures  formed  by  the  chains  of  the  ro- 
settes, may  be  caused  by  the  chemical  action  described  taking  place 
around  the  edges  of  large  masses  or  concretions  of  the  iron  ore. 

These  crystalline  forms  occur  only  in  the  dryer  parts  of  the 
cave.  Where  there  is  more  moisture,  as  in  the  '  Snow-ball  room,' 
the  gypsum  merely  forms  white,  rounded  concretions,  originating 
from  nodules  of  the  iron  ore  on  the  roof  and  sides  of  the  cave.'" 

With  these  general  remarks  on  the  cave  we  give  a  brief  account 
of  its  interesting  fauna,*  comprising  representatives  of  the  Fishes, 
Insects,  and  Crustaceans.  No  Mollusks  nor  Radiates  have  as  yet 
been  discovered,  but  the  lower  forms  of  life  have  been  detected 
by  Tellkampf,  who  collected  several  species  belonging  to  the  gen- 
era Monas,  Chilomonas,  and  (?)  Chilodon. 

*In  the  following  pages  it  will  be  noticed  that  the  authors  have  expressed  widely 
different  views  as  to  the  origin  of  the  peculiar  forms  of  subterranean  animals. 


CHAPTER   II. 

THE   CRUSTACEANS   AND   INSECTS.* 

BY   A.    8.    PACKARD,    JR. 


Fig. 122. 


REPRESENTATIVES  of  all  the  grand  divisions  of  the  Insects  and 
Crustaceans  have  been  found  in  this  cave,  and  if  no  worms  have 
yet  been  detected,  one  or  more  species  would  undoubtedly  reward 
a  thorough  search. 

We  will  enumerate  what  have  been  found,  beginning  with  the 
higher  forms.  No  Hymenoptera  (bees,  wasps,  and  ants)  or  Lepi- 
doptera  (moths)  are  yet  recorded  as  being  peculiar  to  caves.  The 
Diptera  (flies)  are  represented  by  two  species,  one  of  Anthomyia 
(Fig.  122)  or  a  closely 
allied  genus,  and  the  sec- 
ond belonging  to  the  sin- 
gular and  interesting  ge- 
nus Phora  (Fig.  123). 
The  species  of  Antho- 
myia usually  frequent 
flowers  ;  the  larvae  live  in 
decaying  vegetable  mat- 
ter, or,  like  the  onion 
fly,  attack  healthy  roots  ; 
while  the  maggots  of 
Phora  live  in  decaying 
substances.  It  would  be 

Anthomyia. 

presumptuous  in  the  writ- 
er to  attempt  to  describe  these  forms  without  collections  of  spe- 
cies from  the  neighborhood  of  the  cave,  for  though  like  all  the 
rest  of  the  insects  they  were  found  three  or  four  miles  from  the 
mouth,  yet  they  may  be  found  to  occur  outside  of  its  limits,  as 
the  eyes  and  the  colors  of  the  body  are  as  bright  as  in  other 
species. 

Among  the  beetles,  two  species  were  found  by  Mr.  Cooke.     The 


*From  the  AMERICAN  NATURALIST  for  December,  1871. 


(11) 


12 


INSECTS    OF   THE    CAVE. 


Fig. 124. 


Anoptlialmus  Tellkampjli  of  Erichson,  a  Carabid  (Fig.  124),  and 
Adelops  hirtus  Tellkampf  (Fig.  125)  allied  to  Catops,  one  of  the 
Silphidse  or  burying  beetle  family.  The  Anoptlialmus  is  of  a  pale 
reddish  horn  color,  and  is  totally  blind ;  *  in  the  Adelops,  which 
is  grayish  brown,  there  are  two  pale  spots,  which  may  be  rudi- 
mentary eyes,  as  Tellkampf  and  Erichson  suggest.  No  Hemip- 
tera  (bugs)  have  yet  been  found  either  in  the  caves  of  this  coun- 
try or  Europe.  Two  wingless  grasshoppers  (generally  called 
crickets)  like  the  common  species  found  under  stones  (CeutJio- 
philus  maculatus  Harris) ,  have  been  found  in  our  caves ;  one  is 
the  Hadenoecus  subterraneus  (Fig.  126  nat.  size)  described  by  Mr. 
Scudder,  and  very  abundant  in  Mammoth  Cave.  The  other  spe- 
cies is  C.  stygia  Scudder,  from  Hickman's 
cave,  near  Hickman's  land-  Fig.  123. 
ing,  upon  the  Kentucky  river. 
It  is  closely  allied  to  the 
Mammoth  Cave  species.  Ac- 
cording to  Mr.  Scudder,  the 
specimens  of  C.  stygia  were 
found  by  Mr.  A.  Hyatt  "  in 
the  remotest  corner  of  Hickman's  Cave, 
in  a  sort  of  a  hollow  in  the  rock,  not  par- 
ticularly moist,  but  having  only 
a  sort  of  cave  dampness.  They 
were  found  a  few  hundred  feet 
from  the  sunlight,  living  exclu- 
sively upon  the  walls."  Even 
the  remotest  part  of  that  cave  is 
not  so  gloomy  but  that  some 
sunlight  penetrates  it. 
The  other  species  is  found  both  in  Mammoth  Cave,  and  in  the 
adjoining  White's  Cave.  It  is  found  throughout  the  cave,  and 
most  commonly  (to  quote  Mr.  Scudder)  "about  'Martha's  Vine- 
yard' and  in  the  neighborhood  of  'Richardson's  Spring'  where 
they  were  discovered  jumping  about  with  the  greatest  alacrity 
upon  the  walls,  where  only  they  are  found,  and  even  when  dis- 

*In  Erhardt's  cave,  Montgomery  Co.,  Virginia,  Prof.  Cope  found  "four  or  five  spec- 
imens of  a  new  Anoptlialmus,  the  A.  pusio  of  Horn,  at  a  distance  of  not  more  than  three 
hundred  feet  from  its  mouth.  The  species  is  small,  and  all  were  found  together  under 
a  stone.  Their  movements  were  slow,  in  considerable  contrast  to  the  activity  of  ordi- 
nary Carabidze."  Proc.  Amer.  Phil.  Soc.  1869.  p.  178. 


Phora. 


Fig. 125. 


Anophthalmus  Tellkampfii. 


INSECTS    OF   THE    CAVE.  13 

turbed,  clinging  to  the  ceiling,  upon  which  they  walked  easily; 
they  would  leap  away  from  approaching  footsteps,  but  stop  at  a 
cessation  of  the  noise,  turning  about  and  swaying  their  long  an- 
tennae in  a  most  ludicrous  manner,  in  the  direction  whence  the 
disturbance  had  proceeded ;  the  least  noise  would  increase  their 
tremulousness,  while  they  were  unconcerned  at  distant  motions, 
unaccompanied  by  sound,  even  though  producing  a  sensible  cur- 
rent of  air ;  neither  did  the  light  of  the  lamp  appear  to  disturb 
them ;  their  eyes,  and  those  of  the  succeeding  species  (J2.  stygia) 
are  perfectly  formed  throughout,  and  they  could  apparently  see 
with  ease,  for  they  jump  away  from  the  slowly  approaching  hand, 
so  as  to  necessitate  rapidity  of  motion  in  seizing  them." 

Mr.  Henry  Edwards  has  discovered  a  wingless  grasshopper  in  a 
limestone  cave  at  Collingwood,  Massacre  Bay,  Middle  Island,  New 

Fig.  126. 


Jfadenoecus  subterraneus. 

Zealand.  Says  Mr.  Scudder,  who  has  described  the  species  in  the 
"Proceedings  of  the  Boston  Society  of  Natural  History^'  (Vol. 
xii,  1869,  p.  408)  under  the  name  Hadencecus  Edwardsii^  "the 
cave  is  close  to  the  sea  shore,  and  near  a  very  large  coal  deposit, 
which  occasionally  crops  out  in  the  interior.  The  Hadenceci  were 
rather  numerous,  but  very  difficult  to  catch,  disappearing  in  the 
crevices  of  the  rocks  on  the  approach  of  lights.  They  appeared 
to  be  most  abundant  near  the  streams  of  water  which  percolated 
through  the  rocks."  The  wingless  grasshopper  of  the  European 
caves  is  the  Hadenoecus  palpatus  Scudder,  first  described  by  Sulzer 
under  the  name  Locusta  palpata. 

The  Thysanurous  Neuroptera  are  represented  by  a  species  of 
Machilis,  allied  to  our  common  Macliilis  variabilis  Say,  common  in 
Kentucky  and  the  middle  and  southern  states.  So  far  as  Tell- 


INSECTS    OF   THE    CAVE. 


Fig. 127. 


kampfs  figure  indicates,  it  is  the  same  species  apparently,  as  I 
have  received  numerous  specimens  of  this  widely  distributed  form 
from  Knoxville,  Tennessee,  collected  by  Dr.  Josiah  Curtis. 

It  was  regarded  as  a  crustacean  by  Tellkampf,  and  described 
under  the  name  of  Triura  cavernicola.*  He  mistook  the  labial 
and  maxillary  palpi  for  feet  and  regarded 
the  nine  pairs  of  abdominal  spines  as  feet. 
The  allied  species,  M.  variabilis  Say,  is 
figured  in  vol.  v.  pi.  1,  fig.  8,  9  (see  also 
p.  94  of  vol.  v  of  the  NATURALIST). 

An  interesting  species  of  Campodeaf  of 
which  the  accompanying  cut  (Fig.  127)  is 
a  tolerable  likeness,  though  designed  to  il- 
lustrate another  species  (C.  stapliylinus 
Westw.)  was  discovered  by  Mr.  Cooke. 
Both  the  European  and  our  common  spe- 
cies live  under  stones  in  damp  places,  and 
the  occurrence  of  this  form  in  the  water  is 
quite  remarkable.  The  other  species  are 
blind,  and  I  could  detect  no  eyes  in  the 
Mammoth  Cave  specimen. 

A   small   spider  was   captured  b}r  Mr. 
Cooke,  but  afterwards  lost ;  it  was  brown 
in  color,  and  possibly  distinct   from   the 
Anthrobia  monmouthia  Tellkf.  (Fig.  128)  which  is  an  eyeless  form, 
white  and  very  small,  being  but  half  a  line  in  length.     The  family 

*-Professor  Agassiz  in  his  brief  notice  of  the  Mammoth  Cave  animals,  does  not  criti- 
cise Tellkampf  s  reference  of  this  animal  to  the  Crustacea;  and  so  eminent  an  authority 
upon  the  articulates  as  Schiodte  remarks  that  while  "  Dr.  Tellkampf's  account  affords 
us  no  means  of  forming  any  conclusion  as  to  its  proximate  relations,"  that,  hoAvever, 
it "  appears  to  belong  to  the  order  of  Amphipoda,  and  to  have  a  most  remarkable 
structure."  TellkampPs  figure  of  Machilis  is  entirely  wrong  in  representing  the  labial 
and  maxillary  palpi  as  ending  in  claws,  thus  giving  the  creature  a  crustacean  aspect ; 
and  ndeed  he  describes  them  as  true  feet ! 

t  Campod^a  Cookei  n.  sp.  Closely  allied  to  C.  Americana, but  it  is  much  larger;  the  an- 
tennae are  24-jointed  instead  of  20-jointed  as  in  C,  Americana,  and  reach  to  the  basal 
abdominal  segment,  while  in  C.  Americana  they  reach  only  to  the  second  thoracic;  the 
terminal  joints  are  much  longer  than  in  that  species,  the  penultimate  joint  being  one- 
third  longer.  Last  three  abdominal  segments  unequal  (equal  in  C.  Americana)  the  penul- 
timate very  short,  not  half  as  long  as  the  terminal,  which  is  longer  and  slenderer  than  in 
C.  Americana,  while  the  three  are  much  narrower  in  proportion  to  the  rest  of  the  body 
than  in  the  other  species.  Hind  femora  longer  than  in  C.  Americana.  Entirely  white 
and  pilose.  Length  .25  inch,  the  largest  C.  Americana  being  .15  to  .20  inch.  (Anal  sty- 
lets broken  off.)  Several  specimens  were  seen  by  Mr.  C.  Cooke,  but  only  one  was  cap- 
tured in  a  pool  of  water,  two  or  three  inches  deep,  in  company  with  the  C-  cidotea. 


Campodea. 


INSECTS    OF   THE   CAVE. 


15 


of  Harvest  men  is  represented  by  a  small  white  form,  described  by 
Tellkampf  under  the  name  of  Plialangodes  armata  (Fig.  129)  but 
now  called  Acanthocheir  armata  Lucas.  The  body  alone  is  but  half 
a  line  long,  the  legs  measuring  two  lines.  It  should  be  borne  in 
mind  that  many  of  the  spiders,  as  well  as  the  Thysanura,  live  in 
holes  and  dark  places,  so  that  we  would  naturally  find  them  in 
caves.  So,  also,  with  the  Myriopods,  of  which  a  most  remarkable 

form*  (Figs.  130,  and  130  a 
front  of  head)  was  found  by 
Mr.  Cooke,  three  or  four  miles 
from  the  mouth  of  the  cave.  It 
is  the  only  truly  hairy  species 
known,  an  approach  to  it  being 
found  in  Pseudotremia  Vudii 
Cope.  It  is  blind,  the  other  spe- 
cies of  this  group  which  Profes- 
sor Cope  found  living  in  caves 
having  eyes.  The  long  hairs  ar- 
AnthroUa  monmouthia.  ranged  along  the  back,  seem  to 

suggest  that  they  are  tactile  organs,  and  of  more  use  to  the  Thous- 
and legs  in  making  its  way  about  the  nooks  and  crannies  of  a  per- 
petually dark  cave  than  eyes  would  be.  It  was  found  by  Mr. 
Cooke  under  a  stone. 

Prof.  Cope  has  contributed  to  the  "  Proceedings  of  the  American 
Philosophical  Society"  (1869,  p.  171)  an  interesting  account  of  the 

*  Spirostrephon  (Pseudotremia')  Copei  n.  sp.  Head  with  rather  short,  dense  hairs;  no 
eyes,  and  no  ocular  depression  behind  the  antennae,  the  surface  of  the  epicranium  being 
well  rounded  to  the  antennal  sockets;  behind  the  insertion  of  the  antennas  the  sides  of 
the  head  are  much  more  swollen  than  in  S.  lactarius.  Antennae  slender,  with  short 
thick  hairs ;  relative  length  of  joints,  the  6th  being  longest ;  6th,  4th,  5th,  3d,  8th,  7th,  1st, 
the  7th  joint  being  much  thicker  than  the  8th.  Twenty-eight  segments  besides  the  head ; 
they  are  entirely  smooth,  striated  neither  longitudinally  nor  transversely;  a  few  of  the 
anterior  segments  rapidly  decrease  in  diameter  towards  the  head.  The  segments  are 
but  slightly  convex,  and  on  each  side  is  a  shoulder,  bearing  three  tubercles  in  a  trans- 
verse row,  each  giving  rise  to  a  long  stiff  hair  one-half  to  two-thirds  as  long  as  the  seg- 
ment is  thick;  these  hairs  stand  up  thickly  all  over  the  back,  and  may  serve  at  once  to 
distinguish  the  species.  No  pores.  Feet  long  and  slender,  nearly  as  long  as  the  an- 
tennae, being  very  slender  towards  the  claws.  Entirely  white.  Length  of  body  .35 
inch  ;  thickness  .04  inch. 

It  is  nearly  allied  to  Pseudotremia  Vudii  of  Cope.  It  will  be  noticed  that  Professor 
Cope  characterizes  the  genus  Spirostrephon  as  having  "  no  pores  ";  though  we  find  it 
difficult  to  reconcile  this  statement  with  that  of  Wood  who  describes  S.  lactarius  as 
having  "  lateral  pores."  Cope  separates  Pseudotremia  from  Spirostrephon  for  the  rea- 
son that  the  segments  have  "two  pores  on  each  side  the  median  line."  The  present 
species  has  no  pores,  but  seems  in  other  characters  to  be  a  true  Spirostrephon,  and  we 
are  thus  led  to  doubt  whether  Pseudotremia  is  a  well  founded  genus. 


16 


INSECTS    OF   THE    CAVE. 


cave  mammals,  articulates  and  shells  -of  the  middle  states.  He 
says  that  "  myriopods  are  the  only  articulates  which  can  be 
readily  found  in  the  remote  regions  of  the  caves  [of  West  Vir- 
ginia] and  they  are  not  very  common  in  a  living  state."  The 
Pseudotremia  cavemarum  which  he  describes,  "  inhabits  the  deep- 
Fig.  129. 


Acanthocheir  armatct. 

est  recesses  of  the  numerous  caves  which  abound  in  Southern  Vir- 
ginia, as  far  as  human  steps  can  penetrate.  I  have  not  seen  it 
near  their  mouths,  though  its  eyes  are  not  undeveloped,  or  smaller 
than  those  of  many  living  in  the  forest.  Judging  from  its  remains, 
which  one  finds  under  stones,  it  is  an  abundant  species,  though 

Fig.  130.  Fig.  130  a. 


Spirostrephon  Copei. 

rarely  seen  by  the  dim  light  of  a  candle  even  after  considerable 
search.  Five  specimens  only  were  procured  from  about  a  dozen 
caves."  Thev second  species,  P.  Vudii  Cope,  was  found  in  Mont- 
gomery Co.  and  he  thinks  it  was  not  found  in  a  cave.  Professor 
Hyatt  informs  me  that  he  saw  near  the  "Bottomless  Pit"  in  Mam- 


CRUSTACEANS    OF    THE    CAVE. 


17 


moth  Cave,  a  brownish  centipede-like  myriopod,  over  an  inch  in 

length,  which  moved  off  in  a  rapid  zigzag  motion.    Unfortunately 

he  did  not  capture  it. 

Next  to  the  blind  fish,  the  blind  crawfish  attracts  the  attention 

of  visitors  to  the  cave.      This  is  the   Cambarus  pellucidus  (Fig. 
F.; .  Tii .  131,   from  Hagen's 

monograph  of  the 
North  American 
Astacidae)  first  de- 
scribed by  Dr.  Tell- 
k  a  m  p  f .  He  re- 
marks that  "the 
eyes  are  rudiment- 
ary in  the  adults, 
but  are  larger  in 
the  young."  We 
might  add  that  this 
is  an  evidence  that 
the  embryo  devel- 
ops like  those  of 
the  other  species ; 
and  that  the  inher- 
itance of  the  blind 
condition  is  proba- 
bly due  to  causes 
first  acting  on  the  a- 
dults  and  transmit- 
ted to  their  young, 
until  the  production 
of  offspring  that  be- 
come blind  becomes 
a  habit.  This  is 
a  partial  proof  at 
least  that  the  char- 
acters separating 
the  genera  and  spe- 

Cambanis  peUucidus.  c[es  Qf  animals    are 

those  inherited  from  adults,  modified  by  their  physical  surround- 
ings an  1  adaptations  to  changing  conditions  of  life,  inducing  cer- 
tain alterations  in  parts  which  have  been  transmitted  with  more  or 

MAMMOTH   CAVE.  2 


18 


CRUSTACEANS    OF   THE    CAVE. 


Fig. 132. 


Csccidotea  styyia  (side  view). 


Fig.  133. 


less  rapidity,  and  become  finally  fixed  and  habitual.  Prof.  Hagen 
has  seen  a  female  of  Cambarus  Bartonii  from  Mammoth  Cave, 
"  with  the  eyes  well  developed,"  and  a  specimen  was  also  found 
by  Mr.  Cooke.  Prof.  Hagen  remarks  that  "  C.  pellucidus  is  the 
most  aberrant  species  of  the  genus.  The  eyes  are  atrophied, 
smaller  at  the  base,  conical,  instead  of  cylindrical  and  elongated, 
as  in  the  other  species.  The  cornea  exists,  but  is  small,  circular, 
and  not  faceted ;  the  optic  fibres  and  the  dark-colored  pigments 
surrounding  them  in  all  other  spe- 
cies are  not  developed."  It  seems 
difficult  for  one  to  imagine  that  our 
blind  craw  fish  was  created  sud- 
denly, without  the  intervention  of 
secondary  laws,  for  there  are  the 
eyes  more  perfect  in  the  young  than 
the  adult,  thus  pointing  back  to  an- 
cestors unlike  the  species  now  ex- 
isting. We  can  now  understand, 
why  embryologists  are  anxiously 
studying  the  embryology  of  animals 
to  see  what  organs  or  characteristics 
are  inherited,  and  what  originate  de 
novo,  thus  building  up  genealogies, 
and  forming  almost  a  new  depart- 
ment of  science  :  comparative  em- 
bryology in  its  truest  and  widest 
sense. 

Of  all  the  animals  found  in  caves, 
either  in  this  country  or  Europe, 
perhaps  the  most  strange  and  unex- 
pected is  the  little  creature  of  which 
we  now  speak.  It  is  an  Isopod  crus- 
tacean, of  which  the  pill  bugs  or  sow  bugs  arc  examples.  A  true 
species  of  pill  bug  (  Titanethes  albus  Schiodte)  inhabits  the  caves  of 
Carniolia,  and  it  is  easy  to  believe  that  one  of  the  numerous  species 
of  this  group  may  have  become  isolated  in  these  caves  and  modi- 
fied into  its  present  form.  So  also  with  the  blind  Niphargus  sty- 
gius  of  Europe,  allied  to  the  fresh  water  Gammarus  so  abundant 
in  pools  of  fresh  water.  We  can  also  imagine  how  a  species  of 
Asellus,  a  fresh  water  Isopod,  could  represent  the  Idoteidse  in  our 


CiEcidotea  stygia  (dorsal  view). 


CRUSTACEANS    OF    THE    CAVE.  19 

caves,  and  one  may  yet  be  found ;  but  how-  the  present  form  be- 
came a  cave  dweller  is  difficult  of  explanation,  as  its  nearest  allies 
are  certain  species  of  Idotea  which  are  all  marine,  with  the  excep- 
tion of  two  species  :  I.  entomon,  living  in  the  sea  and  also  in  the 
depths  of  the  Swedish  lakes,  as  discovered  by  Loven,  the  distin- 
guished Swedish  naturalist,  while  a  species  representing  this  has 
been  detected  by  Dr.  Stimpson  at  the  bottom  of  Lake  Michigan. 
Our  cave  dweller  is  nearly  allied  to  Idotea,  but  differs  in  being 
blind,  and  in  other  particulars,  and  may  be  called  Ccecidotea  sty- 
giob.*  (Fig.  132  side  view,  enlarged  ;  Fig.  133  dorsal  view  ;  6,  in- 
ner antenna ;  c,  1st  leg.)  It  was  found  creeping  over  the  fine 
sandy  bottom,  in  company  with  the  Campodea,  in  a  shallow  pool 
of  water  four  or  five  miles  from  the  mouth  of  the  cave. 

This  closes  our  list  of  known  articulates  from  this  and  other 
caves  in  this  country,  the  result  of  slight  explorations  by  a  few  in- 
dividuals. The  number  will  be  doubtless  increased  by  future  re- 
search. It  is  to  be  hoped  that  our  western  naturalists  will  thor- 
oughly explore  all  the  sinks  and  holes  in  the  cave  country  of  the 
western  and  middle  states.  The  subject  is  one  of  the  highest  in- 
terest in  a  zoological  point  of  view,  and  from  the  light  it  throws  on 
the  doctrine  of  evolution.  Professor  Schiodte,  the  eminent  Danish 
zoologist,  has  given  us  the  most  extended  account  of  the  cave 
fauna  of  Europe,  which  has  been  translated  from  the  Danish  into 
the  Transactions  of  tke  Entomological  Society  of  London  (new 
series  vol.  1,  1851). 

He  examined  four  caves  ;  namely,  that  of  Adelsberg,  the  Mag- 
dalena  and  Luege  caves,  all  in  the  neighborhood  of  Adelsberg, 

*  Generic  characters.  Head  large,  much  thicker  than  the  body,  and  as  long  as  broad; 
subcylindrical,  rounded  in  front.  No  eyes.  First  antennae  slender,  8-jointed  (2d  anten- 
noe  broken  off).  Abdominal  segments  consolidated  into  one  piece.  Differs  chiefly  from 
Idotea,  to  which  it  is  otherwise  closely  allied,  by  the  8-jointed  (instead  of  4-jointed)  1st 
(inner)  antenna?,  the  very  large  head,  and  by  the  absence  of  any  traces  of  the  three  ba- 
sal segments  of  the  abdomen  usually  present  in  Idotea. 

Specific  chai-acters.  Body  smooth,  pure  white :  tegument  thin,  the  viscera  appearing 
through.  Head  as  wide  as  succeeding  segment,  and  a  little  more  than  twice  as  long. 
Inner  antennne  minute,  slender,  the  four  basal  joints  of  nearly  equal  length,  though  the 
fourth  is  a  little  smaller  than  the  basal  three,  remaining  four  joints  much  smaller  than 
others,  being  one-half  as  thick  and  two-thirds  as  long  as  either  ot  the  four  basal  joints ; 
ends  of  last  four  joints  a  little  swollen,  giving  rise  to  two  or  three  hairs;  terminal  joint 
ending  in  a  more  distinct  knob,  and  bearing  five  hairs.  Segment  of  thorax  very  dis- 
.  tinct,  sutures  deeply  incised;  edges  of  segments  pilose;  abdomen  flat  above,  rounded 
behind,  with  a  very  slight  median  projection ;  the  entire  pair  of  gills  do  not  reach  to 
the  end  of  the  abdomen,  and  the 'inner  edges  diverge  posteriorly.  Legs  long  and  slen- 
der, 1st  pair  shorter,  but  no  smaller  than  the  second.  Length  .25  inch. 


20  GENERAL  REMARKS  ON  CAVE  FAUNA. 

and  the  Corneale  cave  at  Trieste.  The  only  plant  found  was  a 
sort  of  fungus,  Byssus  fulvus  Linn.  The  only  vertebrate  is  the 
singular  salamander,  Hypochthon  (Proteus)  anguinus,  found  in  the 
Magdalina  river.  No  shells  were  found.  Regarding  the  articu- 
lates he  writes : 

"On  searching  along  the  wralls  within  the  entrance  of  the  caves, 
among  the  rubbish  and  the  vegetable  debris  along  the  sides  of  the 
river,  we  meet  wTith  a  considerable  number  of  Insecta,  Myriopoda, 
Arachnida  and  Crustacea,  of  various  families  which  shun  daylight ; 
being  such  species  only  as  inhabit  promiscuously  other  places, 
provided  they  are  moist  and  feebly  illumined.  We  find  species  of 
Pterostichus,  Pristonychus,  Amara,  Quedius,  Homalota,  Omalium, 
Hister,  Trichopteryx,  Cryptophagus,  Atomaria,  Ptinns,  Ceraphron, 
Belyta,  a  grasshopper  of  the  Locust  family,  probably  the  Raphido- 
pJwra  cavicola  Fischer,  as  it  was  only  seen  in  the  larva  state,  Trich- 
optera,  Sciara,  Psychoda,  Phora,  Heteromyza,  Sapromyza,  Tomoce- 
rus,  Linyphia,  Gamasus,  Cryptops,  Julus,  and  Asellus.  In  pro- 
portion as  we  recede  from  the  entrance  the  number  of  species  as 
well  as  individuals  greatly  decreases,  and  at  the  distance  which 
entirely  excludes  the  light,  only  single  individuals  are  found.  In 
the  deepest  recesses  these  species  are  entirely  wanting,  except 
some  few  which  have  been  transported  by  the  current ;  only  a  few 
Diptera  are  found  ;  namely,  a  species  of  Phora,  very  near  P.  (ma- 
culata  Meig.,  Heteromyza  flampes  Zett.,  and  Sapromyza  cJirysoph- 
tlialma  Zett.,  extending  also  very  far  into  the  caves,  even  to  the 
remotest  accessible  places  in  Adelsberg  cave,  more  than  half  an 
hour's  walk  from  its  entrance.  Dead  moths%are  occasionally  found 
far  in  the  caves,  being  left  there  by 'the  bats;  and  likewise  acci- 
dental specimens  of  the  parasites  of  the  latter.  Of  the  five  ear- 
lier known  animals  which  inhabit  these  caves,  I  found  Pnstonyclia 
elegans  Dej.  rather  frequently,  and  Homalota  spelcea  Er.  in  consid- 
erable numbers.  Besides  these  are  Anopihalmus  Sclimidtii,  which 
is  very  rare,  and  the  wood  louse,  Titanethes  alba.  The  new  forms 
he  found  were  a  beetle  (Bathyscia  byssina)  allied  to  our  Adelops  ;* 
Stagdbius  troglodytes,  an  aberrant  genus  of  Silphids  ;  a  Podurid, 
Anurophorus  Stillicidii;  and  the  two  blind  arachnidans,  one  a  spi- 
der allied  to  Dysdera,  the  Stalita  tcenaria,  and  a  false-spider,  Blo- 
thrus  spelceus.  Among  the  Crustacea  he  found  Nipliargus  stygius^ 

*Lud\vig  Muller  enumerates  four  other  species  of  Adelops  from  theee  caves,  and 
three  species  from  France,  and  Maclicerites  spelcetis,  in  Verhandl.  Zool.  Bot.  Vereins, 
Wien,  1855,  p.  505.  See  also  Heller's  Beitrage  zur  Bsterreieh.  Grotteu-Fauna.  (Myrio- 
poda and  Crustacea.)  Vienna,  1858.  He  describes  a  myriopod  with  rudimentary  eyes 
(Trachysphceria  Schmidtii)  allied  to  Glomeris,  and  another  blind  species  (Brachydesmus 
subterraneus)  allied  to  Polydesmus ;  also  a  new  Titlianethes  ( T.  graniyer),  and  notices 
Monolistra  cceca  Gerst.  Wankel  (18(51)  also  found  a  new  rhalangid  (Leiobunum  troglo- 
dytes) witli  distinct  eyes  and  four  species  of  mites  in  the  caves  of  Eastern  Austria.  The 
mites  are  Scyphius  spelceus,  Linopodes  subterraneus,  Gamasus  loricatus  and  G.  niveus. 
Mill  an  additional  species  of  Trachysphama  (T.  HyrtHi).  See  also  Ehrenberg's  list  01 
cave  insects  (Monabsberichte  der  Akad.  Berlin.  18(51.) 

t  Several  species  of  Niphargus  occur  in  the  wells  and  hot  springs  in  Europe.   Accord- 


GENERAL    REMARKS    ON    CAVE    FAUNA.  21 

allied  to  G-ammarus,  which  lives  in  small  pools  of  water  and  is 
white  and  blind  ;  and  the  cave  pill  bug,  Titanethes  albus  (Koch.)." 

.     In  conclusion  Schiodte  remarks  that :  — 

"We  may  with  propriety  apply  the  collective  term  Subter- 
ranean Fauna  to  those  animals  which  exclusively  inhabit  caves, 
and  are  expressly  constructed  for  such  habitations.  Still  there 
is  nothing  in  this  name  which  would  indicate  that  these  animals 
have  any  claim  to  be  considered  as  a  separate  group,  beyond  the 
mere  peculiarity  of  their  common  place  of  abode.  While  a  few 
of  them  possess  such  an  extraordinary  structure  as  to  stand  in  no 
comparison  with  those  animals  which  inhabit  the  light,  there  are 
others,  forming  only  more  characteristic  links  in  the  groups  of 
animals  more  or  less  shy  of  light,  of  which  many  are  found  common 
in  the  localities  of  the  caves  ;  and  some  belong  to  genera  having  a 
wide  local,  as  well  as  geographical,  extension.  We  are  accordingly 
prevented  from  considering  the  entire  phenomenon  in  any  other 
light  than  something  purely  local,  and  the  similarity  which  is  ex- 
hibited in  a  few  forms  (Anophthalmus,  Adelops,  Bathyscia)  be- 
tween the  Mammoth  Cave  and  the  caves  in  Carniola,  otherwise 
than  as  a  very  plain  expression  of  that  analogy,  which  subsists 
generally  between  the  fauna  of  Europe  and  of  North  America. 
Besides,  it  is  clear  to  me  that  the  fauna  of  the  caves  of  Carniola 
consists  of  two  divisions,  of  which  the  essential  character  is  refer- 
able on  the  one  hand  to  the  dark  locality,  and  on  the  other  to  the 
additional  confinement  to  staiactitic  formations  ;  as  yet  we  are  not 

ing  to  Bate  and  Westwood  (British  Sessile  eyed  Crustacea)  •'  the  British  examples  have 
been  obtained  from  artificially  excavated  wells  connected  with  houses  for  domestic 
purposes.  In  some  instances  the  wells  have  been  old,  in  others  but  recently  dug.  In 
their  geological  condition  the  habitats  have  been  equally  various.  At  Corsham  the  well 
exists  in  the  Oolite  formation,  at  Ring-wood  in  chalk-flint  gravel,  at  Mannamead  in 
tlate.  At  Corsham  and  Mannamead  they  are  found  on  a  hill,  at  Ringwood  they  lie  low. 
The  appearance  of  some  of  these  animals  in  a  well  soon  after  its  being  excavated, 
raises  a  question  of  considerable  interest.  Thus  they  were  found  at  upper  Claft'ord,  near 
Andover  and  at  Mannamead,  near  Plymouth,  but  not  a  trace  of  them  was  to  be  found  in 
the  Hurroimding  streams;  in  fact  they  perish  in  the  light.  It  is  impossible  to  regard 
them  as  an  extreme  variety,  or  modification  of  our  only  fresh  water  Aniphipod,  Gam- 
marusfluviaiilus,  since  various  parts  not  only  differ  in  form,  but  some  are  altered  in  char- 
acter; for  example,  the  extraordinary  elongation  and  slenderness  of  one  of  the  branches 
of  each  of  the  last  pair  of  caudal  appendages  seem  to  be  a  special  structure,  having 
for  its  object  the  antenna-like  use  of  a  delicate  apparatus  at  the  extremity  of  the  body. 
....  Although  we  can  find  no  freshwater  ally  to  this  genus  in  the  rivers  and 
streams  of  Europe,  yet  Bruzelius  has  taken  in  the  deep  sea,  near  Bohusia,  a  form  which 
he  has  described  under  the  name  Eriopis  elongata,  approximating  so  nearly  to  it  that  it 
appears  to  be  scarcely  genetically  distinct. 


anus  Koch,  the  embryology  of  which  has  been  studied  by  V.  St.  George)  N.fontanu» 
Bate,  N.  Kochianun  Bate.  Another  generic  form  is  Crangonyx  founded  by  Bate,  which 
also  belongs  to  the  subterranean  fauna.  "  A  single  species  as  yet  is  all  that  has  been 
found  in  England;  but  we  have  little  doubt  but  that  Gammarus  Ermamii  of  Mime  Ed- 
wards which  was  found  by  M.  Ermann  in  the  warm  springs  of  Kamtschatka  belongs 
also  to  this  genus.  It,  is  curious  that  we  should  have  to  record  that  while  the  animals 
of  this  genus,  as  in  the  p:-e:-e .ling(.Niphargtis)  inhabit  the  deep  artificial  wells,  without 
being  known  to  exist  in  our  rivers  and  streams,  its  nearest  allied  form  is  to  be  found 
in  a  marine  genus,  Gammarella." 


22  GENERAL  REMARKS  ON  CAVE  FAUNA. 

able  vigorously  to  discriminate  between  the  two.  We  shall  ac- 
cordingly look  upon  the  subterranean  fauna,  or  more  properly 
faunas,  as  small  ramifications  which  have  penetrated  into  the  earth 
from  the  geographically-limited  faunas  of  the  adjacent  regions ; 
and  which,  as  they  extended  themselves  into  darkness,  have  been 
accommodated  to  surrounding  circumstances.  Animals  not  far  re- 
mote from  the  ordinary  forms,  prepare  the  transition  from  light  to 
darkness.  Next  follow  those  that  are  constructed  for  twilight ; 
and  last  of  all  those  destined  for  total  darkness,  and  whose  struc- 
ture is  quite  peculiar.  Among  these  some  are  adapted  for  special 
localities,  those  which  inhabit  dry  localities  or  detached  little 
reservoirs  being  totally  blind,  while  others,  destined  for  running 
streams,  have  eyes  of  imperfect  construction,  so  as  to  receive  the 
impression  of  rays  of  light,  but  no  proper  image  of  illuminated 
objects.  We  may  therefore  with  tolerable  precision  arrange  the 
inhabitants  of  caverns  under  the  following  heads  :  — 

Shade  animals.  —  Extensive  genera  and  species  inhabiting  cav- 
erns near  their  entrance,  and,  generally,  all  cool,  shady  and  moist 
localities.  Of  these,  those  that  fly  occasionally  enter  far  into  the 
caverns  (Diptera). 

Twilight  animals.  —  They  belong  to  widely  spread  genera,  but 
are  peculiar  to  the  caves,  and  distinguished  by  their  small  eyes. 
They  are  principally  found  near  the  entrances  to  the  caves,  but 
proceed  deeper  into  the  darkness  than  the  shade-animals,  and 
although  wingless,  they  penetrate  often  the  whole  extent  of  the 
dark  space.  —  (Pristonychus  elegans,  Homalota  spelwa.) 

Cave  animals. — The^y  form,  at  least  in  part,  peculiar  genera,  are 
wingless  and  colorless,  as  far  as  the  consistency  of  their  integu- 
ments will  admit,  and  exist  exclusively  in  total  darkness.  The 
terrestrial  division  is  blind  ;  the  aquatic  has  a  perception  of  light. 
To  this  group  belong  all  the  animals  in  the  Mammoth  Cave,  and 
among  those  of  the  caves  of  Carniola,  Auoplithalmus,  Bathyscia, 
perhaps  likewise  Anurophorus  and  Hypochthon,  which,  however, 
may  belong  to  the  following  group. 

Stalactite  cave  animals.  —  Insects,  Arachnidans  and  Crustaceans 
appertaining  to  peculiar  genera,  wingless,  blind,  brightly  colored 
according  to  the  nature  of  their  integuments,  either  light  brown, 
yellowish  white,  or  snow  white,  perhaps  according  to  the  pre- 
ponderance of  the  chitine  ;  living  in  total  darkness,  peculiar  to 
stalactite  caves,  in  part  occupying  the  columns  and  constructed 
accordingly,  either  for  ascent  or  hovering  over  them.  Here  belong 
most  of  the  animals  treated  of  in  this  memoir — Stagobius,  Blo- 
thrus,  Stalita,  Niphargus,  and  Titanethes."  * 

A  pertinent  question  arises  as  to  the  time  of  the  formation  of 
these  caves  and  when  Lhey  became  inhabitable.  As  previously  stat- 

*In  a  note  appended  he  adds  to  the  list  "a  new  cave  crustacean,  Pnlomnn  anopthal- 
mus  Kollar,  said  to  serve  as  food  for  Hypochthon  [the  Salamander],  of  which  last  geuus 
he  discriminates  six  species." 


ORIGIN    OF   CAVE    LIFE.  23 

ed,  the  caves  of  the  western  and  middle  States  are  in  lower  Car- 
boniferous limestone  rocks,  though  the  Port  Kenned}^  cave  explored 
by  Wheatley  and  Copef  is  in  the  Potsdam  limestone.  They  could 
not  have  been  formed  under  water,  but  when  the  land  was  drained 
by  large  rivers.  This  could  not  have  occurred  previous  to  the  Tri- 
assic  period.  Prof.  Dana  in  his  "Manual  of  Geology"  shows  that 
the  Triassic  continent  spread  westward  from  the  Atlantic  coast  "to 
Kansas,  and  southward  to  Alabama ;  for  through  this  great  area 
there  are  no  rocks  more  recent  than  the  Palaeozoic."  "Through  the 
Mesozoic  period  [comprising  the  Triassic,  Jurassic,  and  Creta- 
ceous periods]  North  America  was  in  general  dry  land,  and  on  the 
east  it  stood  a  large  part  of  the  time  above  its  present  level." 
Though  at  the  close  of  these  periods  there  was  a  general  extinc- 
tion of  life,  yet  this  was  not  probably  a  sudden  (one  of  months 
and  even  years),  but  rather  a  secular  extinction,  and  there  may  be 
plants  and  animals  now  living  on  dry  land,  which  are  the  lineal 
descendants  of  mesozoic  and  more  remotely  of  Carboniferous  forms 
of  life.  So  our  cave  animals  may  possibly  be  the  survivors  of  Mes- 
ozoic forms  of  life,  just  as  we  find  'now  living  at  great  depths  in 
the  sea  remnants  of  Cretaceous  life.  But  from  the  recent  explora- 
tions in  the  caves  of  Europe  and  this  country,  especially  the  Port 
Kennedy  cave,  with  its  remarkable  assemblage  of  vertebrates  and 
.insects,  we  are  led  to  believe  from  the  array  of  facts  presented  by 
Prof.  Cope  that  our  true  subterranean  fauna  probably  does  not 
date  farther  back  than  the  beginning  of  the  Quaternary,  or  Post 
pliocene,  period.  We  quote  his  "general  observations"  in  his 
article  on  the  Port  Kennedy  fauna. 

"The  origin  of  the  caves  which  so  abound  in  the  limestones  of 
the  Alleghany  and  Mississippi  valley  regions,  is  a  subject  of  much 
interest.  Their  galleries  measure  many  thousands  of  miles,  and 
their  number  is  legion.  The  writer  has  examined  twenty-five,  in 
more  or  less  detail,  in  Virginia  and  Tennessee,  and  can  add  his 
testimony  to  the  belief  that  they  have  been  formed  by  currents  of 
running  water.  They  generally  extend  in  a  direction  parallel  to 
the  strike  of  the  strata,  and  have  their  greatest  diameter  in  the 
direction  of  the  dip.  Their  depth  is  determined  in  some  measure 
by  the  softness  of  the  stratum,  whose  removal  has  given  them 
existence,  but  in  thinly  stratified  or  soft  material,  the  roofs  or  large 

t  A  notice  of  the  animals  found  in  this  cave  will  be  found  in  the  Proceedings  of  the 
American  Philosophical  Society,  April,  1871.  The  insects  there  enumerated  would 
probably  not  come  under  the  head  of  cave  insects . 


24  ORIGIN    OF    CAVE    LIFE. 

masses  of  rocks  fall  in,  which  interrupt  the  passage  below.  Caves, 
however,  exist  when  the  strata  are  horizontal.  Their  course  is 
changed  by  joints  or  faults,  into  which  the  excavating  waters  have 
found  their  way. 

That  these  caves  were  formed  prior  to  the  postpliocene  fauna  is 
evident  from  the  fact  that  they  contain  its  remains.  That  they 
Were  not  in  existence  prior  to  the  drift  is  probable,  from  the  fact 
that  they  contain  no  remains  of  life  of  any  earlier  period  so  far  as 
known,  though  in  only  two  cases,  in  Virginia  and  Pennsylvania, 
have  they  been  examined  to  the  bottom.  No  agency  is  at  hand  to 
account  for  their  excavation,  comparable  in  potency  and  efficiency 
to  the  floods  supposed  to  have  marked  the  close  of  the  glacial 
period,  and  which  Prof.  Dana  ascribes  to  the  Cham  plain  epoch. 
An  extraordinary  number  of  rapidly  flowing  waters  must  have 
operated  over  a  great  part  of  the  Southern  States,  some  of  them 
at  an  elevation  of  fifteen  hundred  feet  and  over  (perhaps  two  thou- 
sand) above  the  present  level  of  the  sea.  A  cave  in  the  Gap 
Mountain,  on  the  Kanawha  river,  which  I  explored  for  three  miles, 
has  at  least  that  elevation. 

That  a  territory  experiencing  such  conditions  was  suitable  for 
the  occupation  of  such  a  fauna  as  the  deposits  contained  in  these 
caves  reveal,  is  not  probable..  The  material  in  which  the  bones 
occur  in  the  south  is  an  impure  limestone,  being  mixed  with  and 
colored  by  the  red  soil  which  covers  the  surface  of  the  ground.  It 
is  rather  soft  but  hardens  on  exposure  to  the  air. 

The  question  then  remains  so  far  unanswered  as  to  whether  a 
submergence  occurred  subsequent  to  the  development  of  the  post- 
pliocene  mammalian  fauna.  That  some  important  change  took 
place  is  rendered  probable  by  the  fact,  that  nearly  all  the  neotropi- 
cal types  of  the  animals  have  been  banished  from  our  territory, 
and  the  greater  part  of  the  species  of  all  types  have  become  ex- 
tinct. Two  facts  have  come  under  my  observation  which  indicate 
a  subsequent  submergence.  A  series  of  caves  or  portions  of  a 
single  cave  once  existing  on  the  southeast  side  of  a  range  of  low 
hills  among  the  Alleghany  mountains  in  Wythe  Co.,  Virginia,  was 
found  to  have  been  removed  by  denudation,  fragments  of  the  bot- 
tom deposit  only  remaining  in  fissures  and  concavities,,  separated 
by  various  intervals  from  each  other.  These  fragments  yielded  the 
remains  of  twenty  species  of  postpliocene  mammalia.*  This  de- 
nudation can  be  ascribed  to  local  causes,  following  a  subsidence 
of  uncertain  extent.  In  a  cave  examined  in  Tennessee  the  Qssife- 
rous  deposit  was  in  part  attached  to  the  roof  of  the  chamber. 
Identical  fossils  were  taken  from  the  floor.  This  might,  however, 
be  accounted  for  on  local  grounds.  The  islands  of  the  eastern 
part  of  the  West  Indies  appear  to  have  been  separated  by  submer- 
gence of  larger  areas,  at  the  close  of  the  period  during  which  they 

*See  Proceed.  Amer.  Phil.  Soc.  1869, 171. 


ORIGIN    OF    CAVE    LIFE.  25 

were  inhabited  by  postpliocene  mammalia  and  shells.  The  caves 
of  Anguilla  include  remains  of  twelve  vertebrates,*  of  which  seven 
are  mammalia  of  extinct  species,  and  several  of  them  are  of  large 
size.  These  are  associated  with  two  recent  species  of  molluscs 
Turbo  pica,  and  a  Tudora  near  pup&formis.'f  As  these  large  ani- 
mals no  doubt  required  a  more  extended  territory  for  their  support 
than  that  represented  by  the  small  island  Anguilla,  there  is  eA^ery 
probability  that  the  separation  of  these  islands  took  place  at  a 
late  period  of  time  and  probably  subsequent  to  the  spread  of  the 
postpliocene  fauna  over  North  America." 

I  think  the  reader  will  conclude  from  the  facts  Prof.  Cope  so 
clearly  presents,  that  the  subterranean  fauna  of  this  country  does 
not  date  back  of  the  Quaternary  period.  These  species  must  have 
been  created  and  taken  up  their  abode  in  these  caves  (Mammoth 
Cave  and  those  of  Montgomery  Count}r,  Virginia)  after  the  breccia 
flooring  their  bottoms  and  containing  the  bones  of  Quaternary  ani- 
mals had  been  deposited ;  or  else  migrated  from  Tertiary  caves 
farther  south,  which  is  not  probable,  as  it  has  been  previously 
shown  that  those  blind  animals  inhabiting  wells  immediately  die 
on  being  exposed  to  the  light  (British  Sessile-eyed  Crustacea,  i, 
p.  313),  though  the  blind  craw  fish  is  not  thus  affected. 

The  case  becomes  much  simpler  when  we  consider  the  age  of 
the  rocks  in  which  the  Adelsberg  and  other  caves  mentioned  by 
Schiodte  are  situated.  The  Alps  were  under  water  in  the  Middle 
Eocene  ;  consequently  the  caves  could  not  have  been  formed  until 
the  close  of  the  Tertiary.  Hence  the  species  of  the  cave  fauna 
were  evidently  created  either  at  the  close  of  the  Tertiary,  or  more 
probably  the  beginning  of  the  Quaternary,  as  "even  in  the  later 
part  of  the  Pliocene  era  there  was  an  elevation  of  three  thousand 
feet  in  a  part  of  the  Island  of  Sicily"  (Dana).  We  are  therefore 
led  to  conclude  that  the  species  of  the  subterranean  fauna  the 
world  over  are  recent  creations,  probably  not  older  than  the  ex- 
tinct mammals  associated  with  man. 

*Loc.  cit.  1839, 183;  1870,  608.  A  fourth  species  of  gigantic  Chinchillid  has  been  found 
by  Dr.  Rijgersma,  which  may  be  called  Loxomylus  quadrans  Cope.  It  is  represented  by 
portions  of  jaws  and  teeth  of  three  individuals.  It  is  one  of  the  largest  species,  equal- 
ling the  L.  latidens,  and  has  several  marked  characters.  Thus  the  roots  of  the  molars 
are  very  short,  and  the  triturating  surface  oblique  to  the  shaft.  The  roots  of  the  second 
and  fourth  are  longer  than  those  of  the  first  and  third.  The  last  molar  has  four  dental 
columns  instead  of  three  as  in  the  other  Loxomyli,  and  is  triangular  or  quadrant-shaped 
in  section ;  the  third  is  quadrangular  in  section,  and  has  three  columns.  The  second  is 
the  smallest,  being  only  .(>  the  length  of  the  subtriangular,  first.  Length  of  dental  series 
m  .063  or  2.5  inches.  Palate  narrow  and  deeply  concave.  There  is  but  little  or  no  lat- 
eral constriction  in  the  outlines  of  the  teeth;  the  shanks  are  entirely  straight.  In  its 
additional  dentinal  column,  this  species  approaches  the  genus  Amblyrhiza. 

The  large  Chinchillas  of  Anguilla  are  as  follows,  Loxomylus  lortgide 
quadrant* ,  and  AmUi/rhiza  inundata. 

fSee  Bland,  Proceed.  Amer.  Phil.  Soc.,  1871,  58. 


26  ORIGIN    OF    CAVE    LIFE. 

Assuming  on  the  principles  of  evolution  that  the  cave  animals 
were  derived  from  other  species  changed  by  migration  from  the 
outer  world  to  the  new  and  strange  regions  of  total  darkness,  it 
seems  evident  that  geologically  speaking  the  species  were  suddenly 
formed,  though  the  changes  may  not  have  been  wrought  Until  after 
several  thousand  generations.  According  to  the  doctrine  of  natu- 
ral selection,  by  which  species  pass  from  one  into  another  by 
a  great  number  of  minute  variations,  this  time  was  not  sufficient  for 
the  production  of  even  a  species,  to  say  nothing  of  a  genus.  But 
the  comparatively  sudden  creation  of  these  cave  animals  affords,  it 
seems  to  us,  a  very  strong  argument  for  the  theory  of  Cope  and 
Hyatt  of  creation  by  acceleration  and  retardation,  xwhich  has  been 
fully  set  forth  in  this  journal.  The  strongly  marked  characters 
which  separate  these  animals  from  their  allies  in  the  sunlight,  are 
just  those  fitting  them  for  their  cave  life  and  those  which  we  would 
imagine  would  be  the  first  to  be  acquired  by  them  on  being  re- 
moved from  their  normal  habitat. 

On  introducing  the  wingless  locust,  Ceuthophilus  maculatus, 
into  a  cave,  where  it  must  live  not  under  stones,  but  by  clinging  to 
the  walls,  its  legs  would  tend  to  grow  longer,  its  antennae  and 
palpi  would  elongate  and  become  more  delicate  organs  of  hearing 
as  well  as  touch,*  and  the  body  would  bleach  partially  out,  as  we 
find  to  be  the  case  in  H.  subterranea  and  C.  stygia.  The  Carabid 
beetle,  Anopthalmus,  extending  farther  into  the  cave,  would  lose 
its  wings  (all  cave  insects  except  the  Dipterahave  no  wings,  elytra 
excepted)  and  eyes,  but  as  nearly  all  the  family  are  retiring  in 
their  habits,  the  species  hiding  under  stones,  its  form  would  not 
undergo  farther  striking  modification.  So  with  the  blind  Campo- 
dea,  which  does  not  differ  from  its  blind  congeners,  which  live 
more  or  less  in  the  twilight,  except  in  its  antennae  becoming 
longer.  The  blind  Adelops,  but  with  rudiments  of  eyes,  does  not 
greatly  depart  in  habits  from  Catops,  while  on  the  other  hand  the 
remarkable  Stagobius  of  the  Illyrian  caves,  which  according  to 

*  After  writing  this  article,  and  without  knowledge  of  his  views,  we  turned  to  Darwin's 
Origin  of  Species  to  learn  what  he  had  to  say  on  the  origin  of  cave  animals.  He  attri 
butes  their  loss  of  sight  to  disuse,  and  remarks :—"  By  the  time  an  animal  has  reached, 
after  numberless  generations,  the  deepest  recesses,  disuse  will  on  this  view  have  more 
or  less  perfectly  obliterated  its  eyes,  and  natural  selection  will  often  have  effected 
other  changes,  such  as  an  increase  in  the  length  of  the  antenna?  or  palpi,  as  a  compen- 
sation for  blindness."  5th  Amer.  Edit.,  p.  143.  We  are  glad  to  find  our  views  as  to  the 
increase  in  the  length  of  the  antennae  and  palpi  compensating  for  the  loss  of  eyesight, 
confirmed  by  Mr.  Darwin. 


ORIGIN   OF    CAVE    LIFE.  27 

Schiodte  spends  its  life  in  crawling  ten  to  twenty  feet  above  the 
floor  -over  the  columns  formed  by  the  stalactites,  to  which  unique 
mode  of  life  it  is  throughout  perfectly  adapted,  is  remarkably 
different  from  other  Silphids.  Its  legs  are  very  long  and  inserted 
far  apart  (the  prothorax  being  remarkably  long),  with  surprisingly 
long  claws,  while  the  antennae,  again,  are  of  great  length  and 
densely  clothed  with  hairs,  making  them  most  delicate  sense  or- 
gans.* So  also  are  the  limbs  of  the  false  scorpion,  and  the  spi- 
der and  pill  bug  (Titanethes)  of  remarkable  length. 

But  the  modifications  in  the  body  of  the  Spirostrephon  are  such 
that  many  might  deem  its  aberrant  characters  as  of  generic  impor- 
tance. It  loses  its  eyes,  which  its  nearest  allies  in  other,  but 
smaller,  caves  possess,  and  instead  gains  in  the  delicate  hairs  on 
its  back,  which  evidently  form  tactile  organs  of  great  delicacy ; 
the  feet  are  remarkably  long,  as  also  the  antennae.  These  are  not 
new  formations  but  simply  modifications,  apparently  by  use  or  dis- 
use, of  organs  present  in  the  other  species.  The  aberrant  myrio- 
pod  and  Stagobius  are  paralleled  by  the  blind  fish,  an  animal  so 
difficult  to  classify,  and  so  evidently  adapted  for  its  abode  in  end- 
less darkness.  And  as  an  additional  proof  of  the  view  here  taken 
that  these  cave  animals  are  modified  from  more  or  less  allied  spe- 
cies existing  outside  of  the  caves,  we  have  the  case  of  the  craw 
fish,  whose  eyes  (like  those  of  the  mole),  are  larger  in  the  young 
than  adult,  indicating  its  descent  from  a  species  endowed  with  the 
faculty  of  sight,  while  in  the  adult  the  appendages  are  modified  as 
tactile  organs  so  as  to  make  up  for  its  loss  of  eyesight,  in  order 
that  it  may  still  take  its  prey. 

We  thus  see  that  these  cave  animals  are  modified  in  various 
w^iys,  some  being  blind,  others  very  hairy,  others  with  long  ap- 
pendages. All  are  not  modified  in  the  same  way  in  homologous 
organs  ;  another  argument  in  proof  of  their  descent  from  ancestors 

*  Schiodte  remarks  that  "it  is  difficult  to  understand  the  mode  of  life  of  Stagobius 
troglodytes;  or  how  this  slow  and  defenceless  animal  can  escape  being  devoured  by  the 
rapid,  piratical  Arachnidans,  or  find  adequate  support  on  columns,  for  inhabiting 
which  it  is  so  manifestly  constructed.  We  are  led  in  this  respect  to  consider  the  anten- 
na?. Whatever  signilicauce  we  attach  to  those  enigmatical  organs,  we  must  admit  that 
they  are  organs  of  sense,  in  which  view  an  animal  having  them  so  much  developed  as 
Stagobius,  must  possess  a  great  advantage  over  its  enemies,  if  these  be  only  Arachni- 
dans.  Its  cautious  and  slow  progress,  and  its  timid  reconnoitring  demeanor,  fully 
indicate  that  it  is  conscious  of  life  being  in  perpetual  danger,  and  that  it  endeavors  to 
the  utmost  to  avoid  that  danger.  Darkness,  which  always  favors  the  pursued  more 
than  the  pursuer,  comes  to  its  aid,  especially  on  the  uneven  excavated  surface  of  the 
columns." 


ORIGIN    OF    CAVE    LIFE. 

whose  habits  varied,  as  those  of  their  out-of-door  allies  do  at  pres- 
ent. Had  they  been  specially  created  for  subterranean  life,  we 
should  have  expected  a  much  greater  uniformity  in  the  organs 
adapting  them  to  a  cave  life  than  we  actually  find  to  be  the  case. 

Another  fact  of  interest  in  this  connection  is  the  circumstance 
that  these  cave  species  breed  slowly,  being  remarkably  poor  in  in- 
dividuals ;  they  are  nearly  all  extremely  rare.*  Did  they  breed  as 
numerously  as  their  allies  in  the  outer  world  the  whole  race  would 
probably  starve,  as  the  supply  of  food  even  for  those  which  do 
live  is  wonderfully  limited. 

It  is  now  known  that  animals  inhabiting  the  abysses  of  the  sea  are 
often  highly  colored  :  light  must  penetrate  there,  for  we  know  that 
were  the  darkness  total  they  would  be  colorless  like  the  cave  insects. 

In  view  of  the  many  important  questions  which  arise  in  relation 
to  cave  animals,  and  which  have  been  too  imperfectly  discussed 
here,  we  trust  naturalists  the  world  over  will  be  led  to  explore 
caves  with  new  zeal,  and  record  their  discoveries  with  minuteness, 
and  the  greatest  possible  regard  to  exactness.  The  caves  of  the 
West  Indian  Islands  should  first  of  all  be  carefully  explored. 
Also  those  of  Brazil,  those  of  the  East  Indies  and  of  Africa, 
while  fresh  and  most  extended  explorations  of  our  own  Mammoth 
Cave  should  be  made,  perhaps  by  a  commission  acting  under  gov- 
ernment or  State  authority,  in  order  that  the  most  ample  facili- 
ties may  be  afforded  by  the  parties  owning  the  cave. 


NOTE.— Since  my  article  was  printed,  Prof.  Cope's  article  entitled  "Life  in  the  Wyan- 
dotte  Cave"  has  appeared  in  the  "Annals  and  Magazine  of  Natural  History"  (Lon- 
don) for  November.  He  enumerates  the  following  articulates  as  inhabitants  of  this 
cave;  " Anophtlialmus  Telllcampfii,  and  another  species;  two  species  of  Staphylinidae; 
Raphidophora ;  two  species  of  flies;  an  Aranea-like  and  Opiliolike  spider;  a  species  of 
Pseudotremia;  Cambarus  pellucidus,  an  unknown  aquatic  Crustacean  with  external 
egg  pouches,  and  a  Lernsean  (crustacean)  parasitic  on  the  blind  fish.  Of  these  one 
beetle  (Anophtlialmus),  the  cricket  (Raphidophora),  a  fly,  the  Opilio-like  spider,  the  cen- 
tipede, and  the  blind  crawfish,  are  probably  the  same  as  those  found  in  the  Mammoth 
Cave.  Two  beetles  and  two  crustaceans  are  certainly  different  from  those  of  the 
latter,  and  the  centipedes  are  much  more  numerous.  The  Gammaroid  crustarean 
found  in  the  waters  of  the  Mammoth  Cave,  and  which  is,  no  doubt  in  part,  the  food  of 
the  blind  fish,  we  did  not  find;  but  some  such  species  no  doubt  exists,  as  we  found  an 
abundance  of  a  lively  little  tetradecapod  crustacean  near  the  mouth  of  a  cave  close  by." 

*  The  wingless  grasshoppers  are  common  however,  and  Prof.  Hagen  writes  me  that 
the  cave  insects  in  Europe  are  probably  not  so  rare  as  they  are  thought  to  be  by  natu- 
ralists, since  the  guides  do  not  show  the  best  collecting  places,  wishing  to  keep  a  stock 
on  hand  to  sell  to  visitors. 


CHAPTER  III. 

THE   BLIND   FISHES   OF   THE   MAMMOTH   CAVE    AND   THEIR 

ALLIES.* 

BY   F.    W.    PUTNAM. 


THE  blind  fish  of  the  Mammoth  Cave  has  from  its  discovery 
been  regarded  with  curiosity  by  all  who  have  heard  of  its  exis- 
tence, while  anatomists  and  physiologists  have  considered  it  as  one 
of  those  singular  animals  whose  special  anatomy  must  be  studied 
in  order  to  understand  correctly  facts  that  have  been  demonstrated 
from  other  sources  ;  and,  in  these  da}Ts  of  the  Darwinian  and  devel- 
opment theories,  the  little  blind  fish  is  called  forth  to  give  its  tes- 
timony, pro  or  con. 

Before  touching  upon  this  point,  however,  we  must  call  attention 
to  the  structure  of  the  fish  and  its  allies,  and  to  others  that  are 
either  partially  or  totally  blind. 

In  the  lancelet  (Branchiostoma)  and  the  hag  (Myxine)  the  eye 
is  described  "as  simple  in  form  as  that  of  a  leach,  consisting  sim- 
ply of  a  skin  follicle  f  coated  by  a  dark  pigment,  which  receives  the 
end  of  a  nerve  from  the  brain."  Such  an  eye  speck  as  this  struc- 
ture gives  would  only  answer  for  the  simple  perception,  of  light.  In 
the  young  J  of  the  lampreys  (Petromyzon)  the  eye  is  very  small  and 

*  From  the  AMERICAN  XATTRALIST  for  January,  1872. 

t  Sae  further  on  where  Prof.  Wyinan  questions  this  structure. 

t  These  young  lampreys  have  been  described  under  the  generic  name  of  Ammocoetes, 
and  it  was  not  until  1S.36,  when  Prof.  Miiller  discovered  the  fact  of  a  metamorphosis  in 
the  lampreys,  that  their  true  position  was  ascertained.  Prof.  Miiller  has  traced  the 
history  of  the  common  European  species  and  shown  that  it  is  three  or  four  years  in 
attaining  its  perfect  form.  With  this  fact  before  us  and  with  the  early  stages  of  the 
Myxinoids  still  unknown,  have  we  not  some  reason  for  suspecting  that  the  Lancelet  may 
yet  prove  to  be  a  larval  form  of  the  Myxinoids,  notwithstanding  that  it  is  said  to  lay 
eggs  ?  Why  should  we  not  suspect  the  existence  in  the  very  lowest  vertebrates  of  some- 

(29) 


30  BLIND    FISHES    IN    GENERAL. 

placed  in  a  fold  of  the  skin  of  the  head,  and  probably  of  little  use, 
as  these  young  remain  buried  in  the  sand ;  but  as  they  attain  ma- 
turity, and,  with  it,  the  parasitic  habits  of  the  adult,  their  eyes  are 
developed  to  a  fair  size,  thus  reversing  the  general  rule  in  the  class. 

In  most  other  fishes  the  eyes  are  developed  to  a  full  and  even 
remarkable  extent  as  to  size  and  perfection  of  sight  in  water. 
In  Anableps,  or  the  so  called  four  eyed  fish  of  the  fresh  waters  of 
Central  and  South  America,  which  belongs  to  a  closely  allied  fam- 
ily with  our  blind  fish,  the  Cyprinodontidve,  the  eyes  are  not  only 
fully  developed,  but  are  divided  into  an  upper  and  lower  portion 
in  such  a  way,  by  an  opaque  horizontal  line,  as  to  give  the  effect 
of  two  pupils,  by  which  the  fish  probably  sees  as  well  when  follow- 
ing its  prey  on  the  surface  with  its  eyes  out  of  water,  as  when 
under  water.  But  it  is  in  the  interesting  family  of  cat  fishes  (Siiu- 
ridce)  that  we  find  the  most  singulaf  arrangement  of  eyes  in  per- 
fect adaptation  to  the  diversified  modes  of  life  of  the  numerous 
species.  In  this  family  the  eyes  assume  nearly  every  possible  mod- 
ification from  partial  and  even  total  blindness  to  perfectly  develop- 
ed eyes,  and  these  organs  are  placed  in  almost  every  conceivable 
position  in  a  fish's  head  ;  from  the  ordinary  large  eyes  on  the  side, 
to  small  ones  on  top  of  the  head,  enabling  the  fish  to  see  only  what 
is  above ;  to  the  oval  eyes  on  the  side,  in  some  just  back  of  the 
mouth,  situated  in  such  a  way  that  the  fish  can  only  see  what  is  in 
close  proximity  to  its  jaws  or  even  below  them.  Many  genera  of 
this  family  found  in  South  America,*  Africa f  and  Asia,!  have  the 
eyes  so  small  and  buried  under  the  skin  or  protected  by  folds  or 
cartilage,  as  evidently  to  be  of  no  more  use  than  simply  to  distin- 
guish light  from  darkness. 

Among  the  most  interesting  forms  of  this  family,  in  this  respect, 
is  the  genus  described  by  Prof.  Cope  under  the  name  of  Gronias 
nigrilabris.  This  fish  is  very  closely  allied  to  our  common  bull 


thing  akin  to  "alternate  generation,"  or  of  larvae  capable  of  reproduction?  Without 
having  any  facts  to  support  such  an  assumption,  except  that,  on  general  principles,  the 
young  of  Myxine  would  probably  be  very  much  like  Branchiostoma,  and  that  its  young 
is  not  known,  while  Branchiostoma  has  only  been  found  in  waters  where  some  species 
of  Myxinoid  exists,  I  think  that  before  the  position  of  the  laucelet  is  firmly  established 
we  must  know  the  embryology  of  the  Myxinoids;  for  should  the  lancelet  prove  not  to 
be  the  young  of  the  Myxinoids,  it  must  necessarily  form  a  distinct  class  of  animals, 
perhaps  as  near  to  the  mollusks  as  to  the  vertebrates. 

*  Pimelodus  cyclopium  of  Humboldt,  Helogenes,  Agoniosus  and  other  genera. 

f  Eatropius  congensis. 

\Ailia,  Shilbiclithys,  Bagroides  and  other  genera. 


THE    PENNSYLVANIA    BLIND    FISH.  31 

pout  or  horned  pout,  and  of  about  the  same  size  (ten  inches  in 
length).  It  was  taken  in  the  Conestoga  river  in  Lancaster  Co., 
Penn.,  where  it  is  "occasionally  caught  by  fishermen  and  is  sup- 
posed to  issue  from  a  subterranean  stream  said  to  traverse  the 
limestone  in  that  part  of  Lancaster  Co..  and  discharge  into  the 
Conestoga."  We  quote  the  following  from  Prof.  Cope's  remarks 
on  the  fish: *- 

"Two  specimens  of  this  fish  present  an  interesting  condition  of 
the  rudimental  eyes.  On  the  left  side  of  both  a  small  perforation 
exists  in  the  corium,  which  is  closed  by  the  epidermis,  representing 
a  rudimental  cornea  ;  on  the  other  the  corium  is  complete.  Here 
the  eyeball  exists  as  a  very  small  cartilaginous  sphere  with  thick 
walls,  concealed  by  the  muscles  and  fibrous  tissue  attached,  and 
filled  by  a  minute  nucleus  of  pigment.  On  the  other  the  sphere  is 
larger  and  thinner  walled,  the  thinnest  portion  adherent  to  the 
corneal  spot  above  mentioned  ;  there  is  a  lining  of  pigment.  It 
is  scarcely  collapsed  in  one,  in  the  other  so  closely  as  to  give  a 
tripodal  section.  Here  we  have  an  interesting  transitional  condi- 
tion in  one  and  the  same  animal,  with  regard  to  a  peculiarity  which 
has  at  the  same  time  physiological  and  systematic  significance, 
and  is  one  of  the  comparatively  few  cases  where  the  physiological 
appropriateness  of  a  generic  modification  can  be  demonstrated.  It 
is  therefore  not  subject  to  the  difficulty  under  which  the  advocates 
of  natural  selection  labor,  when  necessitated  to  explain  a  structure 
as  being  a  step  in  the  advance  towards,  or  in  the  recession  from, 
liny  unknown  modification  needful  to  the  existence  of  the  species. 
In  the  present  case  observation  on  the  species  in  a  state  of  nature 
may  furnish  interesting  results.  In  no  specimen  has  a  trace  of 
anything  representing  the  lens  been  found." 

When  we  remember  that  the  lens  of  the  eye  in  Amblyopsis  has 
been  found,  even  though  the  eye  is  less  developed  in  all  its  parts 
than  in  Gronias,  it  is  probable  that  a  careful  microscopical  exami- 
nation would  show  its  existence  in  this  genus  also. 

It  is  interesting  to  note  that  this  fish  is  black  above  (lighter  on 
the  sides  and  white  below),  notwithstanding  its  supposed  subter- 
ranean habits,  and  that  all  the  other  members  of  the  family  having 
rudimentary  or  covered  eyes  are  also  dark  colored,  while  the  blind 
fishes  of  the  Mammoth  Cave  and  of  the  caves  in  Cuba  are  nearly 
colorless.  This  want  of  color  in  the  latter  fishes  has  been  consid- 
ered as  due  to  their  subterranean  life.  If  this  be  the  cause,  why 
should  the  blind  cat  fishes  retain  the  colors  characteristic  of  the 
other  members  of  the  family  living  in  open  waters  ? 

*  Proceedings  of  the  Academy  of  Natural  Sciences  of  Philadelphia  for  1864,  p.  231. 


THE    BLIND    FISHES    OF    CUBAN    CAVES. 

The  fishes  which  in  a  general  way,  -so  far  as  blindness,  tactile 
sense  and  mode  of  life  are  concerned,  come  the  nearest  to  the  blind 
fishes  of  the  Mammoth  Cave,  are  those  described  by  Prof.  Poey* 
under  the  names  of  Lucifuga  subterraneus-  and  L.  dentatus^ 
These  fishes  having  the  broad,  flattened,  fleshy  head,  with  minute 
cilia,  without  external  eyes,  and  inhabiting  caves  so  similar  in 
structure  to  the  Mammoth  Cave,  make  a  comparison  of  them  with 
the  fishes  of  the  Mammoth  Cave  most  interesting.  This  is  greatly 
enhanced  by  the  fact  that  the  Cuban  fishes  belong  to  a  family  of 
essentially  marine  habit,  quite  far  removed  from  Amblyopsis. 
The  fresh  water  ling  (Lota),  belonging  to  the  same  great  group 
of  fishes  (though  to  a  distinct  family  or  subfamily)  containing  .the 
cod  on  the  one  hand  and  the  Cuban  blind  fish  on  the  other,  is 
probably  the  nearest  fresh  water  relative  of  the  Cuban  fish,  but 

Fig.  1. 


Blind  Fish  (Stygicola  dentatus)  from  Caves  iu  Cuba. 

the  nearest  representative  yet  known  is  the  marine  genus  Brotula, 
one  species  of  which  is  found  in  the  Caribbean  Sea. 

In  the  Cuban  blind  fish  we  find  ciliary  appendages  on  the 
head  and  body  quite  distinctly  developed,  evidently  of  the  same 
character  as  those  of  Amblyopsis  and  answering  the  purpose  of 
tactile  organs.  These  cilia  are  in  the  form  of  small,  but 'plainly 
visible,  protuberances  (reminding  one  of  the  single  fleshy  protu- 
berance over  the  opercular  opening  just  back  of  the  head  in  Ambly- 
opsis). There  are  eight  of  these  on  top  of  the  head  of  a  speci- 
men I  hastily  examined,  received  from  Prof.  Poey  by  the  Museum 
of  Comparative  Zoology,  and  quite  a  number  arranged  in  three 

*Memorias  Sobre  la  Historia  Natural  de  la  Isla  de  Cuba,  por  Felipe  Poey.  Tomo  2, 
pp.  95-llt.  Pis.  9, 10, 11.  Habana,  1856-8. 

fThis  species  was  afterwards  referred  to  the  genus  Stygicola  Gill,  on  account  of  the 
presence  of  palatine  teeth  which  are  wanting  in  the  other  species.  There  are  also  sev- 
eral other  good  characters,  to  judge  from  the  figures  of  the  head,  skull  and  brain  given 
by  Poey,  that  would  warrant  the  reference  of  the  fish  to  a  distinct  genus  from  L.  suiter- 


FIRST   NOTICE    OF   THE    BLIND    FISH.  33 

rows  on  each  side  of  the  body,  showing  that  tactile  sense  is  well 
developed  in  this  fish ;  though  it  is  rather  singular  that  the  barbels 
on  the  jaws,  so  usually  developed  as  organs  of  touch  in  the  cod 
family  and  its  allies,  are  entirely  wanting  in  this  fish. 

The  brain  of  Lucifuga  subterraneus,  as  represented  by  the  figures 
of  Poey,  differs  very  much  from  that  of  L.  dentatus  and  of  Ambly- 
opsis.  In  all,  the  optic  lobes  are  as  largely  developed  as  in  allied 
fishes  provided  with  well  developed  eyes.  In  Lucifuga  subterra- 
neus  the  cerebral  lobes  are  separated  by  quite  a  space  from  the 
round  optic  lobes,  which  are  represented  as  a  little  larger  than 
the  cerebral  lobes,  and  also  of  greater  diameter  than  the  cerebel- 
lum ;  this  latter  being  more  developed  laterally  than  in  either  L. 
dentatus  or  in  Amblyopsis.  The  three  divisions  of  the  brain 
are  represented,  from  a  top  view,  as  nearly  complete  circles 
(without  division  into  right  and  left  lobes),  of  which  that  repre- 
senting the  optic  lobes  is  slightly  the  largest.  In  L.  dentatus  the 
px'ocencephalon  and  the  optic  lobes  are  represented  as  divided  into 
right  and  left  lobes,  as  in  Amblyopsis,  and  the  cerebellum  does 
not  extend  laterally  over  the  medulla  oblongata  as  in  L.  subterra- 
neus,  but,  as  in  Amblyopsis  (PL  1,  fig.  Id),  is  not  so  broad  as  the 
medulla,  and,  projecting  forwards,  covers  a  much  larger  portion  of 
the  optic  lobes  than  is  the  case  in  L.  subterraneus. 

The  Cuban  blind  fish  has  the  body,  cheeks  and  opercular  bones 
covered  with  scales.  As  in  Amblyopsis  the  eyes  exist,  but  are  so 
imbedded  in  the  flesh  of  the  head  as  to  be  of  no  use.  The  out- 
line cut  here  given  (Fig.  1),  copied  from  Poey,  is  very  character- 
istic of  the  form  of  the  fish,  but  does  not  exhibit  the  fleshy  cilia 
or  details  of  scaling. 

The  first  notice  that  I  can  find  of  the  Mammoth  Cave  blind  fish 
is  that  contained  in  the  "Proceedings  of  the  Academy  of  Natural 
Sciences  of  Philadelphia/'  Vol.  1,  page  175,  where  is  recorded  the 
presentation  of  a  specimen  to  the  Academy  by  "W.  T.  Craige,  M. 
D.,  at  the  Meeting  held  on  May  24,  1842,  in  the  following  words  : — 

"A  white,  eyeless  crayfish  (Astacus  Bartoni?)  and  ajmaall white 
fish,  also  eyeless  (presumed  to  belong  to  a  subgenus  of  Silurus), 
both  taken  from  a  small  stream  called  the  'River  Styx'  in  the 
Mammoth  Cave,  Kentucky,  about  two  and  one-half  miles  from  the 
entrance." 

Dr.  DeKay  in  his  "Natural  History  of  New  York,  Fishes,"  page 
187,  published  in  J842,  describes  the  fish,  from  a  poor  specimen  in 

MAMMOTH   CAVE.  3 


34  THE    BLIND    FISHES    OF    MAMMOTH    CAVE. 

the  Cabinet  of  the  Lyceum  of  Natural  History  of  New  York, 
under  the  name  of  Amblyopsis  *  spelceus.^  DeKay's  description  is 
on  the  whole  so  characteristic  of  the  fish  as  to  leave  no  doubt  as 
to  the  species  he  had  before  him,  though  the  statement  that  it  has 
eight  rays  supporting  the  branchiostegal  membrane  (instead  of 
six),  and  that  the  eyes  are  "large"  but  under  the  skin,  must  have 
been  due  to  the  bad  condition  of  his  specimen  and  to  his  taking 
the  fatty  layer  covering  the  minute  eyes  for  the  eyes  themselves, 
as  pointed  out  by  Prof.  Wyman.  Dr.  DeKay  places  the  genus  with 
the  Siluridse  (cat  fishes)  but  at  "the  same  time  questions  its  con- 
nection with  the  family  and  says  that  it  will  probably  form  the 
type  of  a  new  family.  In  1843  Prof.  Jeffries  Wyman  \  gave  an 
account  of  the  dissection  of  a  specimen  in  which  he  could  not  find 
a  trace  of  the  eye  or  of  the  optic  nerve,  probably  owing  to  the 
condition  of  the  specimen,  as  he  afterwards  §  found  the  eye  spots, 
and  made  out  the  structure  of  the  eye.  When  describing  the 
brain,  Prof.  "Wyman  calls  attention  to  the  fact  of  the  optic  lobes 
being  as  well  developed  as  in  allied  fishes  with  well  developed  eyes, 
and  asks  if  this  fact  does  not  indicate  that  the  optic  lobes  are  the 
seat  of  other  functions  as  well  as  that  of  sight.  He  also  calls 
attention  to  the  papillae  on  the  head  as  tactile  organs  furnished 
with  nerves  from  the  fifth  pair. 

Dr.  Theo.  Tellkampf  ||  was  the  first  to  point  out  the  existence  of 
the  rudimentary  eyes  from  dissections  made  by  himself  and  Prof. 
J.  Miiller,  and  to  state  that  they  can  be  detected  in  some  specimens 
as  black  spots  under  the  skin  by  means  of  a  powerful  lens.  Prof. 
Wyman  afterwards  detected  the  eye  through  the  skin  in  several 
specimens.  Dr.  Tellkampf  also  was  the  first?  to  call  attention  to 
the  "  folds  on  the  head,  as  undoubtedly  serving  as  organs  of  touch, 
as  numerous  fine  nerves  lead  from  the  trigeminal  nerve  to  them 
and  to  the  skin  of  the  head  generally." 

It  is  also  to  Dr.  Tellkampf  that  we  are  indebted  for  the  first 
figure  of  the  fish,^[  and  for  figures  illustrating  the  brain,  and  inter- 
nal organs.  The  descriptions  of  the  anatomy  of  the  fish  by  Drs. 

*  Obtuse  vision,    f  Of  a  cave. 

t  Silliman's  Journal,  Vol.  45,  p.  94. 

§  Proceedings  Boston  Soc.  Nat.  Hist.,  Vol.  4,  p.  395.    1853. 

||  Miiller's  Archiv.  fur  Anat.,  1844.  p.  392.  Reprinted  in  the  New  York  Journal  of  Med- 
icine for  July,  1845.  p.  84,  with  pliito. 

IT  The  only  other  figures  of  the  species,  that  I  am  aware  of,  are  the  simple  outlines 
given  in  Poey's  Mem.  de  Cuba,  the  woodcut  in  Wood's  Illustrated  Natural  History  and 
the  cut  in  Tenney's  Zoology.  None  of  these  figures  are  very  satisfactory. 


THE    BLIND    FISH    FAMILY.  35 

Tellkampf  and  Wyman  are  all  that  have  ever  been  written  on  the 
subject  of  any  importance,  with  the  exception  of  the  description 
of  the  eye  by  Dr.  Dalton,  whose  paper,  in  the  "New  York  Medical 
Times,"  vol.  2,  p.  354,  I  have  not  seen.  Prof.  Poey  gives  a  com- 
parison of  portions  of  the  structure  with  that  of  the  Cuban  blind 
fishes. 

Dr.  Tellkampf  proposed  the  name  of  Heteropygii*  for  the  family 
of  which,  at  the  time,  a  single  species  from  the  Mammoth  Cave  was 
the  only  known  representative,  and  makes  a  comparison  of  the  char- 
acters with  those  of  Apliredoderus  Say  anus,  a  fish  found  only  in  the 
fresh  waters  of  the  United  States,  and  belonging  to  the  old  family 
of  Percoids,  but  now  considered  as  representing  a  family  by  itself, 
though  closely  allied  to  the  North  American  breams  (Pomotis),  and 
having  the  anal  opening  under  the  throat  as  in  the  blind  fish. 

Dr.  Storer,f  not  knowing  of  Dr.  Tellkampf 's  paper,  proposed 
the  name  of  Hypsceidce,  for  the  blind  fish,  and  placed  it  between 
the  minnow  and  the  pickerel  families,  in  the  order  of  Malacoptery- 
gian,  or  soft  rayed,  fishes.  According  to  the  system  adopted  by 
Dr.  Giinther,  it  stands  as  closely  allied  to  the  minnows,  Cyprino- 
dontidce  (many  of  which  are  viviparous  and  have  the  single  ovary 
and  general  character  of  the  blind  fish),  and  the  shiners,  Cyprini- 
dm,  of  the  order  of  Physostomi.  Dr.  Tellkampf,  in  discussing  the 
relations  of  the  family,  points  out  its  many  resemblances  to  the 
family  of  Clupesoces,  and  its  differences  from  the  Siluroids,  Cy- 
prinodontes  and  Clupeoids,  with  which  it  has  more  or  less  affinity, 
real  or  supposed.  Prof.  Cope  in  his  paper  on  the  Classification  of 
Fishes  j  places  the  Arnblyopsis  in  the  order  of  Haplomi  with  the 
shore  minnows,  pickerel  and  mud  fish,  and  in  an  article  on  the  Wy- 
andotte  Cave,§  he  says  that  the  Cyprinodontes  (shore  minnows) 
are  its  nearest  allies.  This  arrangement  by  Prof.  Cope  places 
the  Haplomi  between  the  order  containing  the  herrings  and  that 
containing  the  electric  eel  of  South  America,  all  included  with  the 
garpike,  dog  fish  of  the  fresh  waters  (Amia),  cat  fishes,  suckers 
and  eels  proper,  etc.,  etc.,  in  the  division  of  Physostomi  as  limited 
by  him. 

*From  the  advanced  po^iti  >n  of  the  terminus  of  the  intestine  being  so  different  from 
the  position  wliich  it  has  in  ordinary  fishes. 

t  Synopsis  of  the  Fishes  of  North  America,  published  in  1846. 

J  American  Naturalist,  Vol.  5,  p.  579,  1871. 

§  Indianapolis  Daily  Journal  of  September  5,  1871.  Reprinted  in  Ann.  Mag.  Nat. 
Hist.,  Nov.,  1871. 


36  FISHES    WITH    EYES   ALSO    IN   THE    CAVE. 

Prof.  Agassiz  in  1851*  stated  that  the  blind  fish  was  an  aber- 
rant form  of  the  Cyprinodontes. 

Thus  all  those  authors  who  have  expressed  an  opinion  as  to  the 
position  which  the  fish  should  hold  in  the  natural  system  have 
come  to  the  same  conclusions  as  to  the  great  group,  division,  or 
order,  into  which  it  should  be  placed.  For  all  the  terms  used 
above,  when  reduced  to  any  one  system,  bring  Amblyopsis  into 
the  same  general  position  in  the  S3^stem  ;  its  nearest  allies  be- 
ing the  minnows,  pickerels,  shiners  and  herrings ;  and  unless  a 
careful  study  of  its  skeleton  should  prove  to  the  contrary,  we 
must,  from  present  data,  consider  the  family  containing  Amblyop- 
sis as  more  nearly  allied  to  the  Cyprinodontes,  or  our  common 
minnows  having  teeth  on  the  jaws,  than  to  any  other  family,  differ- 
ing from  them  principally  by  the  structure  of  the  several  parts  of 
the  alimentary  canal  and  the  forward  position  of  its  termination. 

I  have  thus  far  mentioned  only  one  species  of  blind  fish  from 
the  cave,  the  Amblyopsis  spelceus.  The  waters  of  the  cave  not  only 
contain  another  species  of  blind  fish,  differing  from  Amblyopsis  in 
several  particulars,  especially  by  its  smaller  size  and  by  being  with- 
out ventral  fins,  which  I  have  identified  as  the  Typliliclithys  subter- 
raneus  of  Dr.  Girard ;  but  also  a  fish  with  well  developed  eyes, 
as  proved  by  the  account  given  by  Dr.  Tellkampf  and  by  the 
drawing  of  a  fish  found  by  Prof.  Wyman,  in  1856,  in  the  stomach 
of  an  Amblyopsis  he  was  dissecting.  In  order  to  call  attention  to 
the  fact  that  fishes  with  eyes  are  at  times,  if  not  always,  in  the 
waters  of  the  cave,  I  have  reproduced  the  drawing  by  Prof.  Wy- 
man on  plate  1,  fig.  13.  It  is  very  much  to  be  regretted  that  the 
specimen  is  not  now  to  be  found,  and  that  it  was  so  much  acted 
on  by  the  gastric  juice  as  to  destroy  all  external  characters  by 
which  it  could  be  identified  from  the  drawing,  which  is  of  about 
natural  size.  Dr.  Tellkampf's  remarks  on  the  fish  with  eyes  are 
as  follows :  — 

"•  Besides  the  colorless  blind-fish,  there  are  also  others  found  in 
the  cave,  which  are  black,  commonly  known  by  the  name  of  'mud- 
fish.' I  saw  a  dark-colored  fish  in  the  water,  but  did  not  succeed 
in  catching  it.  The  latter  are  said  to  have  eyes,  and  are  entirely 
dissimilar  to  the  blind-fish." 

The  name  "mud-fish,"  given  to  this  fish  with  eyes,  and  the  state- 
ment that  it  is  of  a  dark  color,  together  with  the  drawing  by  Prof. 

*SUliman's  Journal,  p.  128. 


HABITS    OF    THE    BLIND    FISH.  37 

Wyman  of  the  fish  found  in  the  stomach  of  the  blind  fish,  showing 
the  position  of  the  dorsal  fin  to  be  the  same  as  in  the  fish  commonly 
called  mud  fish  in  the  fresh  waters  of  the  Middle,  Western  and 
Southern  States,  perhaps,  indicates  the  fish  with  eyes  to  be  a  spe- 
cies of  Melanura.*  This  fish  is  called  mud  fish  from  the  habit  it  has 
of  burying  itself  in  the  mud,  tail  first,  f  to  the  depth  of  two  to  four 
inches,  and  of  remaining  buried  in  the  mud  in  our  western  ditches 
during  a  time  of  drought.  This  habit,  perhaps,  in  a  measure  fits 
it  for  a  subterranean  life.  The  occurrence  of  a  fish  belonging 
to  the  same  family  with  the  blind  fish,  but  with  well  developed 
eyes,  in  the  subterranean  streams  in  Alabama,  as  mentioned  further 
on  and  figured  on  PL  2,  fig.  4,  however,  renders  it  probable  that  the 
cave  fish  with  eyes  may  be  the  same  or  an  allied  species,  and  the 
drawing  by  Prof.  Wyman  would  answer  equally  as  well  for  it. 

The  farct  that  the  Amblyopsis  succeeded  in  catching  a  fish  of, 
probably,  very  rapid  and  darting  movements,  shows  that  the  tactile 
sense  is  well  developed  and  that  the  blind  fish  must  be  very  active 
in  the  pursuit  of  its  prey ;  probably  guided  by  the  movement 
which  the  latter  makes  in  the  water  so  sensibly  influencing  the  del- 
icate tactile  organs  of  the  blind  fish  that  it  is  enabled  -to  follow 
rapidly,  while  the  pursued,  not  having  the  sense  of  touch  so  fully 
developed,  is  constantly  encountering  obstacles  in  the  darkness. 

In  describing  the  habits  of  the  blind  fish  Dr.  Tellkampf  says  : — 

"It  is  found  solitary,  and  is  very  difficult  to  be  caught,  since  it 
requires  the  greatest  caution  to  bring  the  net  beneath  them  with- 
out driving  them  away.  At  the  slightest  motion  of  the  water  they 
dart  off  a  short  distance  and  usually  stop.  Then  is  the  time  to 
follow  them  rapidly  with  a  net  and  lift  them  out  of  water.  They 
are  mostly  found  near  stones  or  rocks  which  lie  upon  the  bottom, 
but  seldom  near  the  surface  of  the  water." 

Prof.  Cope,  in  describing  the  habits  of  the  blind  fish  which  he 

*  Dr.  Gunther  considers  the  genus  Melanura  of  this  country  to  be  synonymous  with 

Umbla  of  Europe.  In  each 
country  only  one  species  has 
been  as  yet  satisfactorily  de- 
scribed. 


t  See   the  interesting  notes 
on  the  habits  of  the  mud  min- 
now, by  Dr.  Abbott  in  Amer- 
ican Naturalist,  Vol.  4,  pages 
Mud  fish  (Melanura  Umi).  107    and    388,    with    figure    of 

the    fish    on    page   385,  which   we   here   reproduce   for    comparison. 


YOUNG    OF    THE    BLIND    FISH. 

obtained  in  a  stream  that  passes  into  the  'Wyandotte  Cave,  though 
he  entered  it  b}^  means  of  a  well  in  the  vicinity  of  the  cave,  says 
that :  — 

"If  these  Amblyopses  be  not  alarmed  they  come  to  the  surface 
to  feed,  and  swim  in  full  sight  like  white  aquatic  ghosts.  They  are 
then  easily  taken  by  the  hand  or  net,  if  perfect  silence  be  pre- 
served, for  they  are  unconscious  of  the  presence  of  an  enemy 
except  through  the  sense  of  hearing.  This  sense  is,  however,  evi- 
dently very  acute,  for  at  any  noise  they  turn  suddenly  downward, 
and  hide  beneath  stones,  etc.,  on  the  bottom.  They  must  take 
much  of  their  food  near  the  surface,  as  the  life  of  the  depths  is 
apparently  very  sparse.  This  habit  is  rendered  easy  by  the  struc- 
ture of  the  fish,  for  the  mouth  is  directed  upwards,  and  the  head  is 
very  flat  above,  thus  allowing  the  mouth  to  be  at  the  surface." 

The  blind  fish  has  a  single  ovary,  in  common  with  several  genera 
of  viviparous  Cyprinodontes.  In  three  female  specimens  of  Am- 
blyopsis  which  I  have  opened,  the  ovary  was  distended  with  large 
eggs,  but  no  signs  of  the  embryo  could  be  traced.  In  these  three 
specimens  it  was  the  right  ovary  that  was  developed,  and  this,  as 
in  the  figure  (Plate  2,  fig.  Ic),  was  by  the  side  of  the  stomach 
and  did  not  extend  beyond  it.  The  number  of  eggs  contained  in 
the  ovary  was  not  far  from  one  hundred  in  the  specimen  figured. 
As  the  embryos  develop,  the  mass  probably  pushes  further 
back  in  the  cavity  and  also  extends  the  abdominal  walls.  That 
the  fish  is  viviparous  is  proved  by  the  statement  made  by  Mr. 
Thompson  before  the  Belfast  Natural  History  Society,*  that  one 
of  the  blind  fishes  from  the  cave,  four  and  a  half  inches  long, 
uwas  put  in  water  as  soon  as  captured,  where  it  gave  birth  to 
nearly  twenty  young,  which  swam  about  for  some  time,  but  soon 
died.  These,  with  the  exception  of  one  or  two,  were  carefully 
preserved,  and  fifteen  of  them  are  now  before  us  [at  the  meeting, 
I  wish  they  were  here]  ,  they  were  each  four  lines  in  length." 

It  is  singular  that  no  mention  is  made  regarding  these  young,  as 
to  the  presence  or  absence  of  eyes,  and,  as  if  it  was  fated  that  this 
important  point  should  remain  unnoticed  as  long  as  possible,  it  is 
equally  singular  that  Dr.  Stein dachner  omitted  to  examine  some 
very  young  specimens  which  he  received  from  a  friend  a  few 
months  since  and  sent  to  the  Vienna  Museum,  where  they  will 
remain  unexamined  until  he  returns  there.  I  saw  the  Doctor  only 

*  Annals  and  Mag.  of  Natural  History,  Vol.  xiii,  pp.  112, 1844. 


GENERAL    REMARKS    ON    THE    BLIND    FISHES.  39 

a  week  after  these,  to  me,  interesting  specimens  had  been  sent 
abroad,  and  he  was  as  grieved  as  I  was  disappointed  at  my  being 
just  too  late  to  take  advantage  of  them.  (See  note  on  p.  52.) 

At  what  time  the  young  are  born  has  never  been  stated,  but  judg- 
ing from  such  data  as  I  can  at  present  command,  I  think  that  it  must 
be  during  the  months  of  September  and  October.  Specimens  col- 
lected during  those  months  would  probably  contain  embryos  in 
various  stages  of  development,  the  examination  of  which  would  un- 
doubtedly lead  to  most  interesting  results.  (See  note  on  p.  52.) 

Prof.  Wyman  has  most  generously  placed  in  my  hands  his  un- 
published notes  and  drawings  of  the  several  dissections  he  has 
made  of  Amblyopsis,  as  well  as  his  specimens  and  dissections. 
Many  of  these  drawings  are  reproduced  on  Plate  1,  and  will,  with 
his  notes  which  I  here  give,  greatly  enhance  the  value  of  this  arti- 
cle, as  his  dissections  have  been  made  with  the  utmost  care,  and 
with  a  patience  and  delicacy  that  only  a  master  hand  attains.  It 
will  therefore  be  understood  that,  in  giving  credit  to  Prof.  Wyman 
in  the  following  pages,  I  refer  to  his  unpublished  notes,  except 
when  the  quotation  is  given  from  a  special  work.  In  quoting  his 
description  of  the  eye  and  ear  from  "  Silliman's  Journal "  I  have 
changed  the  references  so  as  to  refer  to  his  drawings  reproduced 
on  Plate  1,  and  not  to  the  three  cuts  given  in  "Silliman's  Journal," 
though  the  figures  of  the  brain  and  of  the  otolite  were  copied  from 
those  cuts. 

The  largest  specimens  I  have  seen  of  Amblyopsis  are  several 
males  and  females,  each  from  four  to  four  and  a  half  inches  in 
length,  which  seems  to  be  about  as  large  as  the  fish  grows,  though 
Dr.  Giinther  mentions  a  specimen  in  the  British  Museum  of  five 
inches  in  length.  The  largest  specimen  captured  of  late  years  is 
said  to  have  been  taken,  during  the  summer  of  1871,  and  sold  for 
ten  dollars  to  a  person  who  was  so  desirous  of  securing  the  pre- 
cious morsel  that  he  had  it  cooked  for  his  supper.  The  smallest 
specimen  I  have  seen  was  one  and  nine-tenths  inches  in  length. 
The  general  shape  and  character  of  the  fish  is  best  shown  by  the 
figures  on  plates  1  and  2. 

"The  whole  head,  above  and  below,  is  destitute  of  scales,  the 
naked  skin  extending  backwards  on  the  sides  to  the  base  of  the 
pectoral  fins  ;  the  scaly  portion  of  the  body  above  ends  in  a  semi- 
circular edge  covering  the  space  between  the  upper  ends  of  the 
opercula.  The  skin  covering  the  middle  region  of  the  head  is 


40  TACTILE    ORGANS    OF    BLIND    FISH. 

smooth,  but  on  either  side  is  provided  with  numerous  transverse 
and  longitudinal  ridges  (PI.  1,  fig.  7),  which  are,  on  the  whole, 
regularly  arranged.  The  first  row  of  transverse  ridges,  eight  or 
nine  in  number,  begins  between  the  nostrils  and  extends  back- 
wards, diverging  from  the  median  line.  The  third  ridge  is  crossed 
at  its  outer  end  by  a  longitudinal  one,  as  are  also  two  others  farther 
back.  The  second  and  third  rows,  situated,  in  part,  on  the  sides 
and,  in  part,  on  the  under  surface,  are  less  regular  than  the  preced- 
ing. A  fourth,  on  the  borders  of  the  operculum,  is  still  less  Avell 
defined.  The  transverse  are  also  crossed  here  by  longitudinal 
ridges.  About  ten  vertical  ridges,  also  provided  with  papillae,  and 
similar  to  those  on  the  head,  are  visible  on  the  sides  extending  from 
the  pectoral  fins  to  the  tail,  but  are  not  so  well  defined  as  those  on 
the  head.  The  skin  of  the  head  is  of  extreme  delicacy  and  is  cov- 
ered by  a  very  thin,  loose  layer  of  epithelium."— WYMAN. 

"The  larger  ridges  have  between  twenty  and  thirty  papillae, 
many  of  these  having  a  cup-shaped  indentation  at  the  top,  in 
which  a  delicate  filament  is,  in  some  instances,  seen  (PL  1,  fig.  9). 
These  papillae  are  largely  provided  with  nervous  filaments,  and,  as 
is  obvious  from  their  connection  with  branches  of  the  fifth  pair  of 
nerves,  must  be  considered  purely  tactile,  and  the  large  number  of 
them  shows  that  tactile  sensibility  is  probably  very  acute  and  in 
some  measure  compensates  for  the  virtual  absence  of  the  sense  of 
sight.  Plate  1,  fig.  8,  represents  one  of  the  ridges  of  the  bead 
magnified,  showing  the  papillae  of  which  it  is  made  up,  and  figure 
9  shows  three  papillae  still  more  enlarged.  Two  of  these  show  a 
cup-shaped  cavity  at  the^  top,  and  the  short,  slender  filament  al- 
ready mentioned.  The  surface  of  the  papillae  is  covered  with 
loosely  connected  epithelium  cells.  Fig.  10  shows  the  nervous  fil- 
aments distributed  to  the  papillae  :  a,  a  branch  of  the  fifth  pair  of 
nerves  passing  beneath  the  papillary  ridge  and  sending  filaments 
to  each  papilla.  These  papillary  branches  interchange  filaments, 
forming  a  nervous  plexus  in  connection  wTith  each  ridge.  This 
figure  of  the  nerves  was  drawn  with  a  camera  lucida,  from  a  speci- 
men treated  with  acetic  acid." — WYMAN. 

"Plate  1,  fig.  6,  represents  a  double  system  of  subcutaneous  ca- 
nals, which  extend  the  whole  length  of  the  head,  but  were  not 
traced  farther  back  than  the  edge  of  the  naked  or  scaleless  skin 
which  covers  it.  Forwards  they  bifurcate,  nearly  encircling  the 
nasal  cavity,  towards  the  middle  line  ending  in  a  blind  pouch. 


THE    EYES    OF    THE    BLIND    FISHES.  41 

The  lateral  branch  was  not  traced  distinctly  to  an  end,  but  seemed 
to  connect  with  the  olfactory  cavit}\  The  walls  of  these  canals 
are  exceedingly  delicate  and  easily  overlooked." — WYMAN. 

"Plate  1,  fig.  5,  shows  the  globe  of  the  eye  with  the  optic  nerve 
(c),  as  seen  under  the  microscope.  The  lens  (b)  is  detached  from 
its  proper  place  by  the  pressure  of  the  glass.  Irregularly  arranged 
muscular  bands  are  attached  to  the  exterior  of  the  globe  (a,  a,  a, 
a),  but  were  not  recognized  as  the  homologues  of  the  muscles  of 
the  normal  eye  of  fishes  ;  nevertheless,  they  indicate  that  the  globe 
was  movable." — WYMAN. 

"In  the  three  specimens  recently  dissected,  the  eyes  were  ex- 
posed only  after  the  removal  of  the  skin,  and  the  careful  separation 
from  them  of  the  loose  areolar  tissue  which  fills  the  orbit.  In  a 
fish  four  inches  in  length  the  eyes  measured  one-sixteenth  of  an 
inch  in  their  long  diameter,  and  were  of  an  oval  form  and  black.  A, 
filament  of  nerve  (PI.  1,  fig.  3  a)  was  distinctly  traced  from  the 
globe  to  the  cranial  walls,  but  the  condition  of  the  contents  of  the 
cranium,  from  the  effects  of  the  alcohol,  was  such  as  to  render  it 
impracticable  to  ascertain  the  mode  of  connection  of  the  optic 
nerve  with  the  optic  lobes. 

9  Examined  under  the  microscope  with  a  power  of  about  twenty 
diameters,  the  following  parts  were  satisfactorily  made  out  (PI.  1 , 
fig.  3)  :  1st,  externally  an  exceedingly  thin  membrane,  6,  which 
invested  the  whole  surface  of  the  eye  and  appeared  to  be  continu- 
ous with  a  thin  membrane  covering  the  optic  nerve,  and  was 
therefore  regarded  as  a  sclerotic  ;  2d,  a  layer  of  pigment  cells,  d, 
for  the  most  part  of  a  hexagonal  form,  and  which  were  most  abun- 
dant about  the  anterior  part  of  the  eye ;  3d,  beneath  the  pigment 
a  single  layer  of  colorless  cells,  c,  larger  than  a  pigment  cell,  and 
each  cell  having  a  distinct  nucleus  ;  4th,  just  in  front  of  the  globe  ; 
a  lenticular-shaped,  transparent  body,  e  [see  also  fig.  4],  which 
consisted  of  an  external  membrane  containing  numerous  cells  with 
nuclei.  This  lens-shaped  body  seemed  to  be  retained  in  its  place 
by  a  prolongation  forwards  of  the  external  membrane  of  the  globe  ; 
5th,  the  globe  was  invested  by  loose  areolar  tissue,  which  adhered 
to  it  very  generally,  and  in  some  instances  contained  yellow  fatty 
matter  ;  in  one  specimen  it  formed  a  round  spot,  visible  through  the 
skin  on  each  side  of  the  head,  which  had  all  the  appearance  of  a 
small  eye ;  its  true  nature  was  determined  by  the  microscope 
only.  It  is  not  improbable  that  the  appearance  just  referred  to 
may  have  misled  Dr.  DeKay  —  where  he  states  that  the  eye  exists 
of  the  usual  size,  but  covered  by  the  skin. 

If  the  superficial  membrane  above  noticed  is  denominated  cor- 
rectly the  sclerotic,  then  the  pigment  layer  ma^y  be  regarded  as  the 
representation  of  the  choroid.  The  form  as  well  as  the  position 
of  the  transparent  nucleated  cells  within  the  choroid  correspond 


EYES    AND    EARS    OF    THE    BLIND    FISH. 

for  the  most  part  with  the  retina.  All  of  the  parts  just  enumer- 
ated are  such  as  are  ordinarily  developed  from  and  in  connection 
with  the  encephalon,  and  are  not  in  any  way  dependent  upon  the 
skin.  But  if  the  lenticular-shaped  body  is  the  true  representative 
of  the  crystalline  lens,  it  becomes  difficult  to  account  for  its  pres- 
ence in  Ambtyopsis  according  to  the  generally  recognized  mode  of 
its  development  (since  it  is  usually  formed  from  an  involution  of 
the  skin)  unless  we  suppose  that  after  the  folding  in  of  the  skin 
had  taken  place  in  the  embryonic  condition,  the  lens  retreated  from 
the  surface,  and  all  connection  with  the  intagument  ceased. [*] 

According  to  Quatrefages,  however,  the  eye  of  Amphioxus  [|] 
is  contained  wholly  in  the  cavity  of  the  dura  mater,  and  yet  it  has 
all  the  appearance  of  being  provided  with  a  lens.  If  his  descrip- 
tion be  correct,  then  the  mode  of  development  as  well  as  the  mor- 
phology of  the  eye  in  this  remarkable  fish  is  different  from  that  of 
most  other  vertebrates,  since  the  lens  never  could  have  been 
^formed  from  an  involution  of  the  skin,  nor  could  the  eye  with  its 
lens,  as  Prof.  Owen  asserts,  be  a  modified  cutaneous  follicle. 
Whatever  views  be  taken  with  regard  to  the  development  of  the 
eye  of  the  blind  fish,  the  anatomical  characters  which  have  been 
enumerated  show,  that  though  quite  imperfect  as  we  see  it  in  the 
adult,  it  is  constructed  after  the  type  of  the  eyes  of  other  ver- 
tebrates. It  certainly  is  not  adapted  to  the  formation  of  ima- 
ges, since  the  common  integument  and  the  areolar  tissue  which  are 
interposed  between  it  and  the  surface,  would  prevent  the  transmis- 
sion of  light  to  it  except  in  a  diffused  condition.  No  pupil  or  any- 
thing analogous  to  an  iris  was  detected,  unless  we  regard  as  repre- 
senting the  latter  the  increased  number  of  pigment  cells  at  the 
anterior  part  of  the  globe. 

It  is  said  that  the  blind  fishes  are  acutely  sensitive  to  sounds 
as  well  as  to  undulations  produced  by  other  causes  in  the  water. 
In  the  only  instance  in  which  I  have  dissected  the  organ  of  hear- 
ing (which  I  believe  has  not  before  been  noticed),  all  its  parts 
were  largely  developed,  as  will  be  seen  by  reference  to  PI.  1, 
fig.  1  e.  As  regards  the  general  structure,  the  parts  do  not  differ 
materially  from  those  of  other  fishes  except  in  their  proportional 
dimensions.  The  semi-circular  canals  are  of  great  length,  and 
the  two  which  unite  to  enter  the  vestibule  by  a  common  duct,  it 
will  be  seen,  project  upwards  and  inwards  under  the  vault  of  the 
cranium,  so  as  to  approach  quite  near  to  the  corresponding  parts 
of  the  opposite  side.  The  otolite  contained  in  the  utricle  was  not 
remarkable,  but  that  of  the  vestibule  (PL  1,  fig.  2)  and  seen  in 

*In  birds  and  mammals  there  is  a  stage  of  development  where  the  lids  come  to- 
gether and  firmly  unite,  to  separate  again  when  the  animal  <;gets  its  eyes  open."  In 
the  mole  rat  (Spalax  typhlus)  of  Siberia,  the  lids  never  open,  and  the  eyes  remain 
through  life  covered  with  hairy  skin.  It  is  not  improbable  that  in  Amblyopsis  some- 
thing analogous  to  this,  a  closing  of  the  skin  over  the  eye,  may  have  taken  place. — J.  W. 

1 1  have  used  the  prior  name  of  Branchiostoma  in  this  paper  when  speaking  of  the 
Lancelot. 


GEOGRAPHICAL    RANGE    OF    BLIND    FISH.  43 

dotted  outline  in  fig.  1  e  is  quite  large  when  compared  with  that  of 
a  Leuciscus  of  about  the  same  dimensions  as  the  blind-fish  here 
described."  —  WYMAN,  SillitnaiCs  Journal,  Vol.  17,  p.  259,  1854. 

The  Amblyopsis  spelceus  undoubted!}7  has  quite  an  extensive 
distribution,  probably  existing  in  all  the  subterranean  rivers  that 
flow  through  the  great  limestone  region  underlying  the  Carbonif- 
erous rocks  in  the  central  portion  of  the  United  States.  Prof. 
Cope  obtained  specimens  from  the  Wyandotte  Cave  and  from  wells 
in  its  vicinity,  and  in  the  Museum  of  Comparative  Zoology  at 
Cambridge  there  is  a  specimen  labelled  "from  a  well  near  Lost 
River,  Orange  Co.,  Ind.,"  which,  with  those  from  the  Wyandotte 
Cave,  is  conclusive  evidence  of  its  being  found  on  the  northern 
side  of  the  Ohio*  as  well  as  on  the  southern,  in  the  rivers  of  the 
Mammoth  Cave.  I  have  been  able  to  examine  a  number  of  speci- 
mens from  the  Mammoth  Cave,  and  have  carefully  compared  with 
them  the  one  from  the  well  in  Orange  Co.,  Ind.,  and  find  that  the 
specific  characters  are  remarkably  constant. 

In  1859 1  Dr.  Girard  described  a  blind  fish,  received  by  the 
Smithsonian  Institution  from  J.  E.  Younglove,  Esq.,  who  obtained 
it  "from  a  well  near  Bowling  Green,  Ky."  The  general  appear- 
ance of  this  fish,  which  was  only  one  and  a  half  inches  in  length, 
was  that  of  Amblyopsis  spdwus,  but  it  differed  from  that  species 
in  several  characters,  especially  by  the  absence  of  ventral  fins. 
Dr.  Girard  therefore  referred  the  fish  to  a  distinct  genus  under  the 
name  of  Typhlichthys\  subterraneus.  Dr.  Giinther§  considers  this 
fish  a  variety  of  Amblyopsis  spelceus  and  records  the  specimen  in 
the  British  Museum  "'  from  the  Mammoth  Cave,"  as  "  half-grown."  |j 

By  the  kindness  of  Prof.  Agassiz,  I  have  been  enabled  to  exam- 
ine nine  specimens  of  blind  Jish  without  ventrals,  in  the  Museum  of 
Comparative  Zoology.  Seven  of  these  were  collected  in  the  Mam- 
moth Cave  by  Mr.  Alpheus  Hyatt  in  September,  1859.  One  was 
from  Moulton,  Lawrence  County,  Alabama,  presented  by  Mr. 
Thomas  Peters  ;  and  another  from  Lebanon,  Wilson  Co.,  Tennes- 
see ;  presented  by  Mr.  J.  M.  Safford.  It  is  not  stated  whether 

*  I  have  also  been  informed  by  Mr.  Holmes  of  Lansing,  Mich.,  that  blind  fishes  have 
been  drawn  out  of  wells  in  Michigan, 
t  Proceedings  Acacl.  Nat.  Sci.  Philad.,  p.  63. 

J  Blind  fish. 

§  Catalogue  of  Fishes  in  the  British  Museum,  Vol.  7,  p.  2, 1868. 

II  The  largest  specimen  I  have  seen  of  Typhlichthys  is  one  and  seventeen-twentieths 
inches  in  length,  and  the  smallest  Amblyopsis  one  and  eighteen-twentieths  inches. 


44  OTHER    SPECIES    OF    THE    FAMILY. 

these  latter  came  from  wells  or  caves,  but  probably  from  wells. 
They  are  all  of  about  the  same  size,  one  and  one-half  to  two 
inches  in  length,  and  are  constant  in  their  characters.  Moreover, 
four  of  the  seven  specimens  from  the  Mammoth  Cave  were  females 
with  eggs.  These  eggs  were  as  large  in  proportion  as  those  from 
Amblyopsis.  The  ovary  was  single  and  situated  on  the  right 
side  of  the  stomach,  as  in  Amblyopsis.  The  difference  in.  the 
number  of  eggs  was  very  remarkable,  each  of  the  four  specimens 
examined  having  but  about  thirty  eggs  in  the  ovary,  while  in 
three  females  of  Amblyopsis  (all,  however,  of  nearly  three  times 
the  size  of  Typhlichthys)  there  were  about  one  hundred  eggs  in 
each.  As  in  both  species  there  were  no  signs  of  the  embryos  in 
the  eggs,  it  is  not  probable  that  any  of  the  eggs  had  been  developed 
and  the  young  excluded,  nor  is  it  at  all  likely  that  the  great  vari- 
ation in  the  number  of  eggs  would  simply  indicate  different  ages. 
By  a  reference  to  the  figures  (PL  2),  it  will  be  seen  that  the  pyloric 
appendages,  stomach  and  scales  of  the  two  fishes  are  different. 
For  these  reasons,  taken  in  connection  with  the  absence  of  ven- 
tral fins,  I  have  no  hesitation  in  accepting  Dr.  Girard's  name  as 
valid  for  this  genus,  of  which  we  thus  far  know  of  but  one  species, 
with  a  subterranean  range  from  the  waters  of  the  Mammoth  Cave, 
south  to  the  northern  portion  of  Alabama.  In  this  connection  it 
would  be  most  interesting  to  know  the  relations  of  the  "blind 
fishes"  said  to  have  been  found  in  Michigan.  For  thus  far  we 
have  Typhlichthys  limited  to  the  central  and  southern  portion  of 
the  subterranean  region,  Amblyopsis  to  the  central,  and  the  spe- 
cies in  the  northern  portion  undetermined. 

In  1853,  on  his 'return  from  a  tour  through  the  southern  and 
western  states,  Prof.  Agassiz  gave  a  summary  of  some  of  his 
ichthyological  discoveries  in  a  letter  to  Prof.  J.  D.  Dana.*  In  this 
letter  are  the  following  remarks  :  — 

"I  would  mention  foremost  a  new  genus  which  I  shall  call  Cho- 
logaster,  very  similar  in  general  appearance  to  the  blind  fish  of  the 
Mammoth  Cave,  though  provided  with  eyes  ;  it  has,  like  Ambly- 
opsis, the  anal  aperture  far  advanced  under  the  throat,  but  is  en- 
tirely deprived  of  ventral  fins ;  a  very,  strange  and  unexpected 
combination  of  characters.  I  know  but  one  species,  Ch.  cornutus 
Ag.  It  is  a  small  fish  scarcely  three  inches  long,  living  in  the 
ditches  of  the  rice  fields  in  South  Carolina.  I  derive  its  specific 

*  Published  in  American  Journal  of  Sci.  and  Arts,  Vol.  16  (2d  series),  p.  131,  1853. 


OTHER    FISHES    OF    THE    FAMILY    WITH    EYES.  45 

name  from  the  singular  form  of  the  snout,  which  has  two  horn-like 
projections  above." 

This  is  the  only  information  ever  published  regarding  this  inter- 
esting fish  and  the  only  specimens  known  are  those  on  which  Prof. 
Agassiz  based  the  above  remarks. 

By  the  kindness  of  Professor  L.  Agassiz,  who  has  placed  all 
the  specimens  of  the  family  contained  in  the  Museum  of  Com- 
parative Zoology  in  my  hands  for  study,  I  am  enabled  to  give  a 
figure  and  description  of  this  interesting  species  from  the  three 
specimens  in  the  Museum,  which  were  labelled  as  the  originals  of 
Chologaster  cornutus  Ag.,  from  Waccamaw,  S.  C.,  presented  by 
Mr.  P.  C.  J.  Weston,  1853.  The  largest  of  these  specimens  was 
distended  with  eggs  and  I  was  enabled  to  compare  the  ovary  with 
that  of  Amblyopsis.  From  its  being  single  and  the  eggs  very 
large,  I  have  no  doubt  that  it  is  a  viviparous  fish  like  the  other 
genera  of  the  family.  The  position  of  the  ovary  behind  the  stom- 
ach, as  shown  in  fig.  2c,  plate  2,  and  the  presence  of  four  pyloric 
appendages  (PL  2,  fig.  2  a)  instead  of  two,  as  in  Amblyopsis  (fig. 
la)  and  Typhlichthys  (fig.  3 a),  are  good  internal  characters,  sep- 
arating it  from  the  other  genera,  independently  of  the  presence 
of  eyes  and  the  absence  of  ventral  fins  and  papillary  ridges. 

The  stability  of  the  internal  characters  I  have  mentioned  was 
most  unexpectedly  substantiated  by  the  discovery  of  a  second 
species  (PL  2,  figs.  4,  4  a)  of  the  genus  among  the  specimens  in 
the  Museum  of  Comparative  Zoology.  I  have  the  pleasure  of 
dedicating  this  species*  to  Professor  Agassiz,  not  only  in  kindly 
remembrance  of  the  eight  years  I  was  associated  with  him  as  stu- 
dent and  assistant,  but  also  because  the  fish  so  well  illustrates  the 
decided  position  he  has  taken  relative  to  the  immutability  of  spe- 
cies. 

The  only  specimen  known  of  this  second  species  was  drawn 
from  a  well  in  Lebanon,  Tenn.,  and  presented  to  the  Museum  by 
Mr.  J.  M.  Safford,  Jan.,  1854.  It  is  a  more  slender  fish  than  C. 
cornutus,  but  the  intestine  follows  the  same  course  and  the  four 
pyloric  appendages  are  present  as  in  that  species. 

In  the  genus  Chologaster f  we  have  all  the  family  characters  as 
well  expressed  as  in  the  blind  species,  though  it  differs  from  Am- 

*  A  Synopsis  of  this  family  with  descriptions  of  the  four  species  will  appear  in  the 
"  Report  of  the  Peabody  Academy  of  Science  for  1871."    (Reprinted here.  p.  55.) 
t  Literally  "  bile-stomach;"  probably  named  from  the  yellow  color  of  the  fish. 


46  ORIGIN    OF    THE    BLIND    FISHES. 

blyopsis  and  Typhlichthys  by  the  presence  of  eyes,  the  absence  of 
papillary  ridges  on  the  head  and  body,  and  by  the  longer  intestine 
and  double  the  number  of  pyloric  appendages,  as  well  as  by  the 
position  of  the  ovary  ;  and  agrees  with  Typhlichthys  by  the  ab- 
sence of  ventral  fins.  Amblyopsis  and  Typhlichthys  are  nearly 
colorless,  while  Chologaster  Agassizii  is  of  a  brownish  color  similar 
to  many  of  the  minnows,  and  C.  cornutus  is  brownish  j'ellow,  with 
dark,  longitudinal  bands. 

Among  the  most  interesting  points  in  the  history  of  this  genus 
is  the  fact  of  its  occurring  in  two  widely  different  localities,  C. 
Agassizii  having  been  found  in  a  well,  in  the  same  vicinity  (proba- 
bly in  the  same  well)  with  a  specimen  of  Typhlichthys,  and 
undoubtedly  belonging  to  the  same  subterranean  fauna  west  of  the 
Appalachian  ridge,  while  C.  cornutus  belongs  to  the  southern  coast 
fauna  of  the  eastern  side  of  that  mountain  chain,  and  is  thus  far 
the  only  species  of  the  family  known  beyond  the  limits  of  the  great 
subterranean  region  of  the  United  States. 

Having  now  given  an  outline  of  the  structure,  habits  and  distri- 
bution of  the  four  species  belonging  to  the  family,  and  recapitu- 
lated the  known  facts,  we  are  better  able  to  consider  the  bearings 
of  the  peculiar  adaptation  of  the  blind  fishes,  in  the  Mammoth 
and  other  caves,  to  the  circumstances  under  which  they  exist. 

Prof.  Cope  in  stating,  in  his  account  of  the  blind  fish  of  the 
Wyandotte  Cave,  "that  the  projecting  under  jaw  and  upward  di- 
rection of  the  mouth  renders  it  easy  for  the  fish  to  feed  at  the  sur- 
face of  the  water,  where  it  must  obtain  much  of  its  food,"  suggests 
that :  - 

"This  structure  also  probably  explains  the  fact  of  its  being  the 
sole  representative  of  the  fishes  in  subterranean  waters.  No  doubt 
many  other  forms  were  carried  into  the  caverns  since  the  waters 
first  found  their  way  there,  but  most  of  them  were  like  those  of 
our  present  rivers,  deep  water  or  bottom  i'eeders.  Such  fishes 
would  starve  in  a  cave  river,  where  much  of  the  food  is  carried  to 

them  on  the  surface  of  the  stream The  shore  minnows 

are  their  nearest   allies,  and  many  of   them  have  the  upturned 

mouth  and  flat  head Fishes  of  this,  or  a  similar  famity, 

enclosed  in  subterranean  waters  ages  ago,  would  be  more  likely  to 
live  than  those  of  the  other,  and  the  darkness  would  be  very  apt 
to  be  the  cause  of  the  atrophy  of  the  organs  of  sight  seen  in  the 
Amblyopsis." 

This  suggestion  was  undoubtedly  hastily  made  by  Prof.  Cope 
when  writing  the  letter  which  was  printed  in  the  "Indianapolis 


ORIGIN    OF   THE    BLIND    FISHES.  47 

Journal,"  and  were  it  not  tbat  the  article  has  been  reprinted  in  the 
"  Annals  and  Magazine  of  Natural  History,"  I  should  not  criticise 
the  statement  made  in  an  off-hand  letter  for  publication  in  a  news- 
paper ;  for  with  Prof.  Cope's  knowledge  of  fishes  it  could  simply  be 
a  hasty  thought  which  he  put  on  paper,  when  he  suggests  that  it  is 
because  the  Cyprinodontes  have  a  mouth  directed  upwards  and 
are  surface  feeders  that  they  were  better  adapted  to  a  subterranean 
life  than  other  fishes,  and  hence  maintained  an  existence,  while 
other  species,  which  he  supposes  were  introduced  into  the  subter- 
ranean streams  at  the  same  time,  died  out. 

If  the  fishes  of  the  subterranean  streams  came  from  adjoining 
rivers,  why  were  not  many  of  the  Percoids,  Cyprinoids  and  other 
forms,  that  are  as  essentially  surface  feeders  as  the  Cyprinodon- 
tes (many  of  the  latter  are  purely  "mud  feeders"),  as  capable  of 
maintaining  an  existence  in  the  subterranean  waters  as  any  species 
of  the  latter  ?  Neither  is  it  necessary  for  us  to  assume  that  the 
structure  of  the  fish  should  be  adapted  to  feeding  on  the  surface, 
for  not  only  have  we  in  the  blind  cat  fish,  described  by  Prof. 
Cope  himself,  from  the  subterranean  stream  in  Pennsylvania, 
an  example  of  a  fish  belonging  to  an  entirely  different  family  of 
bottom  feeders,  thriving  under  subterranean  conditions,  but  the 
blind  fishes  of  the  Cuban  caves  are  of  the  great  group  of  cod  fishes 
which  are,  with  hardly  an  exception,  bottom  feeders.  The  fact 
that  the  food  of  the  blind  fishes  of  the  Mammoth  Cave  consists  in 
great  part  of  the  cray  fish  found  in  the  waters  of  the  cave,  as 
shown  by  the  contents  of  several  stomachs  I  have  examined,  and 
also  that  one  blind  fish  at  least  made  a  good  meal  of  another  fish, 
as  already  mentioned,  shows  that  the}T  are  not  content  with  simply 
waiting  for  what  is  brought  to  them  on  the  surface  of  the  water, 
and  that  they  are  probably  as  much  bottom  as  surface  feeders. 

Again,  in  regard  to  the  sense  of  sight,  why  is  it  necessary  to 
assume  that  because  fishes  are  living  in  streams  where  there  is  lit- 
tle or  no  light,  that  it  is  the  cause  of  the  non  development  of 
the  eye  and  the  development  of  other  parts  and  organs?  If  this 
be  the  cause,  how  is  it  that  the  Chologaster  from  the  well  in  Ten- 
nessee, or  the  "mud  fish"  of  the  Mammoth  Cave  are  found  with 
eyes?  Why  should  not  the  same  cause  make  them  blind  if  it  made 
the  Amblyopsis  and  Typhlichthys  blind?  Is  not  the  fact,  pointed 
out  by  Prof.  Wyman,  that  the  optic  lobes  are  as  well  developed  in 
Amblyopsis  as  in  allied  fishes  with  perfect  eyes,  and,  I  may  add, 


48  ORIGIN    OF   THE    BLIND    FISHES. 

as  well  developed  as  those  of  Chologaster  cornutus,  an  argument 
in  favor  of  the  theory  that  the  fishes  were  always  blind  and  that 
they  have  not  become  so  from  the  circumstances  under  which  they 
exist?  If  the  latter  were  the  case  and  the  fishes  have  become  blind 
from  the  want  of  use  of  the  eyes,  why  are  not  the  optic  lobes  also 
atrophied,  as  is  known  to  be  the  case  when  other  animals  lose  their 
sight  ?  I  know  that  many  will  answer  at  once  that  Amblyopsis 
and  Typhlichthys  have  gone  on  further  in  the  development  and 
retardation  of  the  characters  best  adapting  them  to  their  subterra- 
nean life,  and  that  Chologaster  is  a  very  interesting  transitionary. 
form  between  the  open  water  Cyprinodontes  and  the  subterranean 
blind  fishes.  But  is  not  this  assumption  answered  by  the  fact  that 
Chologaster  has  every  character  necessary  to  place  it  in  the  same 
family  with  Amblyopsis  and  Typhlichthys,  while  it  is  as  distinctly 
and  widely  removed  from  the  Cyprinodontes  as  are  the  two  blind 
genera  mentioned  ? 

Assuming,  for  the  moment,  that  Chologaster  is  a  transitional 
form  between  the  surface  feeding  Cyprinodontes,  and  Typhlichthys 
and  Amblyopsis,  let  us  recapitulate  the  characters  that  distinguish 
the  different  forms  and  see  if  they  exhibit  transitions,  and  if  Cho- 
logaster is  traversing  the  slow  developmental  road  to  Ambtyopsis. 

Allowing  all  characters  embraced  in  the  general  structure  of  the 
skeleton,  brain,  scales,  fins,  etc.,  as"  ordinal,  and  common  to  both 
Cyprinodontes  and  Heteropygii,  we  will  recapitulate  only  such  as 
can  be  considered  of  family  and  generic  value  in  the  two  groups. 

CYPRINODONTES.    CHOLOGASTER.    TTPHLICHTHYS.    AMBLYOPSIS. 
Surface  feeders.    In  part.  Unknown.  Partially.  The  same. 

The  same. 


The    same. 


Viviparous.  Many  genera.  Probably.  Probably.  Undoubted- 

*y- 

Ovary.  Single  in  vivipa-    Single    and    Single      and    The  same, 

rous  genera*       placed  behind       placed  at  eide 
and   placed  by      the  stomach.         of  stomach, 
the  side  of  in- 
testine in  some 
and  posterior  in 
others. 

*  The  ovary  is  also  single  in  other  genera  of  viviparous  flahes  belonging  to  distinct 
orders. 


Intestine. 

Stomach  tfpylor- 
ic  appendages. 

In   many   genera 
long  and  convo- 
luted, in  others 
short  and  with 
single  turn. 

In    most,    if   not 
all,  stomach  not 
well  defined 
from  intestine 
and  without  ap- 
pendages. 

Moderately 
long  with  two 
turns. 

Stomach      well 
defined,    003- 
cal,  with  two 
p  y  1  o  r  i  c  a  p- 
penrlages    on 
each  side. 

Shorter  with  two 
turns. 

The  same,  with 
one  pyloric  ap- 
pendage  on 
each  side. 

ORIGIN   OF   THE    BLIND    FISHES. 


49 


Anal  opening. 

In    normal    posi- 

Forward of  pec- 

The  same. 

The  same. 

tion. 

torals.f 

Air  bladder. 

Present     in    few 

Present. 

The  same. 

The  same: 

genera.  J 

Scales. 

On     body     regu- 
larly   imbricat- 

Irregularly   ar- 
ranged, firm- 

The same. 

The  same. 

ed  and  loosely 

ly      attached 

attached. 

by  being  cov- 

ered in  great 

part    by    the 

cuticle. 

Head  with  scales 

With  scales. 

Naked. 

The  same-. 

The  same. 

or  naked.  § 

Tactile  papillce  || 

Absent. 

Absent. 

Very  prominent 

The  same. 

on  the  head  and 

as    ridges    on 

body. 

the  head    and 

sides  of  body. 

Central  fins.V 

Present   in    most 

Absent. 

Absent. 

Present. 

genera,    absent 

in  at  least  two. 

Eyes** 

Well  developed  in 
all. 

Well  developed 
and  normal. 

Rudiment  a  ryft 
and  of  no  use. 

The  same. 

Habitat. 

Fresh  water  ; 

Limestone  wa- 

Limestone    wa- 

The same. 

brackish  water; 

ter  of  subter- 

ter of  subter- 

salt water. 

ranean      riv- 

ranean rivers. 

ers.  Brackish 

water  ? 

Geographical 
range. 

Nearly  all  parts  of 
the  world. 

One  species  in 
subterranean 

Central  &  south- 
ern portion  of 

Central  and 
N.  central 

streams  of  S. 

subterranean 

portion  of 

central     por- 

fauna of  Unit- 

same. 

tion  of  the  U. 

ed  States. 

S.;  a  2d  spe- 
cies in  the  So. 

Atl.  coast  fau- 

na of  U.S. 

From  this  brief  comparison  of  some  of  the  prominent  charac- 
ters of  the  genera  of  the  Heteropygii  with  the  Cyprinodontes,  their 

f  Aphredoderus  and  Gymnotus,  and  other  genera  of  distinct  orders  have  this  forward 
position  of  the  anus  also. 

J  The  air  bladder  is  in  several  families  present  in  some  species  and  absent  in  others. 

§  The  presence  or  absence  of  scales  on  the  head,  or  on  portions  of  it,  is  a  generic 
character  subject  to  great  variation  in  many  families  and  quite  constant  in  others. 

II I  cannot  recall  anything  but  the  barbels  on  the  head  and  jaws  of  many  genera  of 
Cyprinoids,  Siluroids,  Gadoids,  etc.,  etc.,  that  can  be  said  to  be  tactile  organs  among 
fishes,  with  the  exception  of  the  fleshy  papillae  on  the  head  and  body  of  the  blind  fishes 
of  the  American  and  Cuban  caves,  and  the  filaments  of  the  fin  rays  of  many  fishes 
and  the  fleshy  ventral  rays  of  the  Gurnards. 

IT  Of  all  fins,  the  ventrals  are  the  most  likely  to  deviate  from  their  normal  structure 
and  position.  Their  presence  or  absence,  as  exhibited  in  many  families,  and  often  by 
different  ages  of  the  same  fish,  and  the  great  variation  in  their  ppsition  in  different 
genera  of  the  same  family,  render  any  change  in  them  of  either  generic,  specific,  or 
individual  character,  or  simply  indicative  of  age  (as  they  are  lost  in  some  adult  fishes 
while  present  in  the  young,  and  in  others  not  developed  until  after  the  other  fins). 

**  As  I  have  alluded  to  the  fact,  in  the  first  part  of  this  paper,  the  eyes  of  fishes  are 
no  more  the  constant  and  unvarying  part  of  the  fish  structure  than  the  ventral  fins,  and 
like  them  are  subject  to  almost  every  conceivable  variation  in  position  in  the  head,  and 
perfection  in  structure. 

tfThe  largest  specimen  I  have  seen  of  Typhlichthys,  is  less  than  two  inches  in  length 
and  as  the  eye  of  an  A^mblyopsis  of  twice  the  size  is  not  over  a  32d  of  an  inch  in  width 
it  must  be  very  small  indeed  in  Typhlichthys,  and  I  confess  to  not  being  able  to  find  it 
in  an  ordinary  dissection,  assisted  only  by  a  good  lens. 
MAMMOTH   CAVE.  4 


50         .  ORIGIN    OF   THE    BLIND    FISHES. 

acknowledged  nearest  allies,  we  can  only  trace  what  could  be 
regarded  as  a  transition,  or  an  acceleration,  or  a  retardation  of 
development,  in  simply  those  very  characters,  of  eyes  and  ventral 
fins,  that  are  in  themselves  of  the  smallest  importance  in  the  struc- 
ture (permanence  of  character  considered)  of  a  fish,  and,  as  if  to 
show  that  they  were  of  no  importance  in  this  connection,  we  find 
in  the  same  cave,  blind  fishes  with  ventrals  and  without ;  and  in 
the  same  subterranean  stream,  a  blind  fish  and  another  species  of 
the  family  with  well  developed  eyes. 

If  it  is  by  acceleration  and  retardation  of  characters  that  the 
Heteropygii  have  been  developed  from  the  Cyprinodontes,  we  have 
indeed  a  most  startling  and  sudden  change  of  the  nervous  s}^stem. 
In  all  fishes  the  fifth  pair  of  nerves  send  branches  to  the  various 
parts  of  the  head,  but  in  the  blind  fishes  these  branches  are  devel- 
'  oped  in  a  most  wonderful  manner,  while  their  subdivisions  take 
new  courses  and  are  brought  through  the  skin,  and  their  free  ends 
become  protected  by  fleshy  papillae,  so  as  to  answer,  by  their  deli- 
cate sense  of  touch,  for  the  absence  of  sight.  At  the  same  time 
the  principle  of  retardation  must  have  been  at  work  and  checked 
the  development  of  the  optic  nerve  and  the  eye,  while  accelera- 
tion has  caused  other  portions  of  the  head  to  grow  and  cover  over 
the  retarded  eye. 

Now,  if  this  was  the  mode  by  which  blindness  was  brought  about 
and  tactile  sense  substituted,  why  is  it  that  we  still  have  Cholo- 
gaster  Agassizii  in  the  same  waters,  living  under  the  same  condi- 
tions, but  with  no  signs  of  any  such  change  in  its  senses  of  sight 
and  touch  ?  It  may  be  said  that  the  Chologaster  did  not  change 
because  it  probably  had  a  chance  to  swim  in  open  waters  and 
therefore  the  eyes  were  of  use  and  did  not  become  atrophied. 
We  can  only  answer,  that  if  the  Chologaster  had  a  chance  for 
open  water,  so  did  the  Typhlichtlrys  and  yet  that  is  blind. 

If  the  Heteropygii  have  been  developed  from  Cyprinodontes, 
how  can  we  account  for  the  whole  intestinal  canal  becoming  so 
singularly  modified,  and  what  is  there  in  the  difference  of  food  or 
of  life  that  would  bring  about  the  change  in  the  intestine,  stomach 
and  pyloric  appendages,  existing  between  Chologaster  and  Typh- 
lichthys  in  the  same  waters  ?  'To  assume,  that  under  the  same  con- 
ditions, one  fish  will  change  in  all  these  parts  and  another  remain 
intact,  by  the  blind  action  of  uncontrolled  natural  laws,  is,  to  me, 
an  assumption  at  variation  with  facts  as  I  understand  them. 


ORIGIN    OF    THE    BLIND    FISHES.  51 

Looking  at  the  case  from  the  standpoint  which  the  facts  force 
me  to  take,  it  seems  to  me  far  more  in  accordance  with  the  laws  of 
nature,  as  I  interpret  them,  to  go  back  to  the  time  when  the 
region  now  occupied  by  the  subterranean  streams,  was  a  salt  and 
brackish  water  estuary,  inhabited  by  marine  forms,  including  the 
brackish  water  forms  of  the  Cyprinodontes  and  their  allies  (but  not 
descendants)  the  Heteropygii.  The  families  and  genera  having  the 
characters  they  now  exhibit,  but  most  likely  more  numerously  rep- 
resented than  now,  as  many  probably  became  exterminated  as  the 
salt  waters  of  the  basin  gradually  became  brackish  and  more  lim- 
ited, as  the  bottom  of  this  basin  was  gradually  elevated,  and 
finalty,  as  the  waters  became  confined  to  still  narrower  limits  and 
changed  from  salt  to  brackish  and  from  brackish  to  fresh,  only 
such  species  would  continue  as  could  survive  the  change,  and  they 
were  of  the  minnow  type  represented  by  the  Heteropygii,  and  per- 
haps some  other  genera  of  brackish  water  forms  that  have  not 
yet  been  discovered. 

In  support  of  this  hypothesis  we  have  one  species  of  the  family, 
Chologaster  cornutus,  now  living  in  the  ditches  of  the  rice  fields  of 
South  Carolina,  under  very  similar  conditions  to  those  under  which 
others  of  the  family  may  have  lived  in  long  preceding  geological 
times  ;  and  to  prove  that  the  development  of  the  family  was  not 
brought  about  by  the  subterranean  conditions  under  which  some 
of  the  species  now  live,  we  have  the  one  with  eyes  living  with  the 
one  without,  and  the  South  Carolina  species  to  show  that  a  sub- 
terranean life  is  not  essential  to  the  development  of  the  singular 
characters  which  the  family  possess. 

That  a  salt  or  brackish  water  fish  would  be  most  likely  to  be 
the  kind  that  would  continue  to  exist  in  the  subterranean  streams, 
is  probable  from  the  fact  that  in  all  limestone  formations  caves 
are  quite  common,  and  would  in  most  instances  be  occupied  first 
with  salt  water  and  then  brackish,  and  finally  with  fresh  water  so 
thoroughly  impregnated  with  lime  as  to  render  it  probable  that 
brackish  water  species  might  easily  adapt  themselves  to  the 
change,  while  a  pure  fresh  water  species  might  not  relish  the  solu- 
tion of  lime  any  more  than  the  solution  of  salt,  and  we  know  how 
few  fishes  there  are  that  can  live  for  even  an  hour  on  beino- 

Of 

changed  from  fresh  to  salt,  or  salt  to  fresh,  water.  We  have  also 
the  case  of  the  Cuban  blind  fishes  belonging  to  genera  with  their 
nearest  representative  in  the  family  a  marine  form,  and  with  the 


52  NOTE    ON   THE   YOUNG    BLIND    FISH. 

whole  family  of  cods  and  their  allies,  to  which  group  they  belong, 
essentially  marine.  Further  than  this  the  cat  fish  from  the  subter- 
ranean stream  in  Pennsylvania  belongs  to  a  family  having  many 
marine  and  brackish  water  representatives.  As  another  very  in- 
teresting fact  in  favor  of  the  theory  that  the  Heteropygii  were 
formerly  of  brackish  water,  we  have  the  important  discovery  by 
Prof.  Cope  of  the  Lernsean  parasite  on  a  specimen  of  Amblyopsis 
from  the  Wyandotte  cave  ;  this  genus  of  parasitic  crustaceans  be- 
ing very  common  on  marine  and  migratory  fishes,  and  much  less 
abundant  on  fresh  water  species. 

Thus  I  think  that  we  have  as  good  reasons  for  the  belief  in 
the  immutability  and  early  origin  of  the  species  of  the  family  of 
Heteropygii,  as  we  have  for  their  mutability  and  late  development, 
and,  to  one  of  my,  perhaps,  too  deeply  rooted  ideas,  a  far  more 
satisfactory  theory  ;  for,  with  our  present  knowledge,  it  is  but  the- 
ory on  either  side. 

YOUNG  OF  THE  BLIND  FISH.  —  Dr.  Hagen  gives  me  the  follow- 
ing information  about  the  young  specimens  I  mentioned  (page  38) 
as  belonging  to  Dr.  Steindachner,  which  I  just  missed  seeing 
before  they  were  sent  to  Vienna.  These  specimens  were  procured 
by  Dr.  Hartung  for  Dr.  Steindachner  under  the  following  circum- 
stances. Just  as  Dr.  Hartung  was  leaving  the  cave  hotel  on  Oct. 
21,  a  bottle  was  brought  to  him  containing  four  specimens,  one  of 
which  was  smaller  than  the  others  (probably  Typhlichthys) ,  all 
living.  He  immediately  transferred  them  to  a  jar  containing  alco- 
hol and  took  no  notice  of  them  until  he  reached  Nashville,  when 
he  discovered  an  addition  of  eight  little  ones  in  the  jar. 

The  birth  of  these  young  was  undoubtedly  due  to  placing  the 
parent  in  the  alcohol,  and  the  date  (Oct.  21)  would  correspond  to 
the  time  I  have  stated  as  probably  that  at  which  the  young  were 
born. 

Dr.  Hagen  states  that  he  examined  the  young  under  a  lens,  with- 
out taking  them  from  the  jar,  and  could  not  discover  any  eyes.  The 
specimens  were  about  three  lines  in  length. 

So  now  we  have  two  more  facts  to  add  to  the  history  of  the 
blind  fishes  (though  whether  they  apply  to  Amblyopsis  or  Typh- 
lichthys is  not  yet  settled).  First,  that  the  young  are  born  in 
October,  and  second,  that  they  are  without  external  eyes  when 
born. — From  the  AMERICAN  NATURALIST  for  February,  1872. 


\  A 


I 


G.A.WALKER-  sc. 


EXPLANATION   OF   PLATE   ONE. 


[All  the  figures  on  this  plate  are  from  original  drawings  by  Prof.  J.  Wyman.] 

FIG.  1.  Brain,  nerves  and  organ  of  hearing  of  Amblyopsis  spelaus;  enlarged;  a,  olfac 
tory  lobes  and  nerves;  6,  cerebral  lobes;  c,  optic  lobes;  d,  cerebellum;  «, 
organ  of  hearing,  showing  the  semicircular  canals,  with  the  otolite  repre- 
sented in  place  by  the  dotted  lines;  /,  medulla  oblongata;  g,  optic  nerves 
and  eye  specks. 

FIG.  2.    Otolite,  enlarged. 

FIG.  3.  Eye,  magnified  (natural  size  one-sixteenth  of  an  inch  in  length) ;  a,  optic  nerve ; 
6,  sclerotic  membrane ;  c,  layer  of  colorless  cells ;  d,  layer  of  pigment  cells 
(iris?);  e,  lens. 

FIG.  4.    Lens,  enlarged  and  showing  the  cells. 

FIG.  5.  Eye,  enlarged,  showing  the  muscular  bands,  a,  a,  a,  a;  b,  the  lens  pressed  out 
of  place;  c,  the  optic  nerve. 

FIG.  6.  Top  of  head,  showing  the  canals  under  the  skin,  of  the  natural  size.  The  two 
black  dots  and  lines  indicate  the  eyes  and  optic  nerves  in  position. 

FIG.  7.    Top  of  head,  showing  the  arrangement  of  the  ridges  of  papillae.    Natural  size. 
FIG.  8.    One  of  the  ridges  of  papilla?  fr*om  the  head,  magnified. 

FIG.  9.  Three  of  the  papilla?  from  the  ridge,  still  more  magnified,  showing  the  cup- 
shaped  summit  and  projecting  filament. 

FIG.  10.  A  portion  of  the  ridge  magnified,  and  treated  with  acid,  to  show  the  arrange- 
ment of  the  nervous  plexus  supplying  the  papillae  with  nerve  filaments  from 
a  branch  (a)  of  the  fifth  pair. 

FIG.  11.  Epithelial  cells  from  the  head. 
FIG.  12.  Epithelial  cells  from  the  body. 
FIG.  13.  A  fish  with  eyes,  found  in  the  stomach  of  an  Amblyopsis. 


(53) 


EXPLANATION   OF  PLATE  TWO. 


FIG.  1.  AMBLYOPSIS  SPEL^EUS  DeKay.    Natural  size. 

la.  Stomach  and  pyloric  appendages.    Twice  natural  size. 

1  &.  Scale,  magnified. 

Ic.  Abdominal  cavity,  showing  position  of  stomach  and  single  ovary.    Natural 
size. 

FIG.  2.  CHOLOGASTER  CORNUTUS  Agassiz.    Natural  size. 

2  a.  Stomach  and  pyloric  appendages.    Twice  natural  size. 

2  &.  Scale,  magnified. 

2c.  Abdominal  cavity,  showing  stomach  and  single  ovary  behind  the  stomach. 
Twice  natural  size. 

FIG.  3.  TYPHLICHTHYS  SUBTERRANETJS  Girard.    Slightly  more  than  natural  size. 

3a.  Stomach  and  pyloric  appendages.    Twice  natural  size. 

3  &.  Scale,  magnified. 


FIG.  4.      fnoT  nc  \  STFT?  AGASSIZII  Putnam.    Natural  size. 

4  a.    Stomach  and  pyloric  appendages.    Twice  natural  size. 
4&.    Scale,  magnified. 

The  scales  figured  on  the  plate  are  all  from  the  second  or  third  row  under  the  dorsal 
fin.  4  b  is  represented  with  the  posterior  margin  dotvn,  all  the  others  are  represented 
with  the  posterior  margin  on  the  left.  The  natural  size  of  the  scales  is  given  by  the 
minute  outline  at  the  left  of  the  figures  above  each  scale  ;  4  &  is  so  small  that  the  natural 
size  can  hardly  be  represented  by  the  black  dot. 


(54) 


PI    2 


TJHIVERSITT 


CHAPTER  IV. 
#t*« 

SYNOPSIS  OF  THE  FAMILY  HfiT-E»e£¥6«: 


BY  F.  "W.  PUTNAM. 


HETEROPYGII  TELLKAMPF,  Muller's  Arch.  f.  Anat.,  p.  392, 
1844;  and  New  York  Journal  of  Medicine,  v,  p.  84,  1845. 

Hypsczidce  Storer,  Synopsis  N.  A.  Fish,  p.  435, 1846. 

Brain  of  ordinary  development  in  all  its  parts,  similar  to  that  of 
Cyprinodontes  and  of  about  the  same  proportions.  Cerebral  lobes 
larger  than  the  nearly  round  optic  lobes.  Cerebellum  overlapping  the 
posterior  third  of  the  optic  lobes.  Medulla  oblongata  broad,  with 
well  defined  right  and  left  sides.  (On  comparing  the  brains  of  the 
three  genera  the  only  difference  noticed  \vas  that  in  Chologaster  the 
cerebellum  was  not  quite  as  large  proportionally,  but  more  elongated 
and  not  quite  as  wide  as  in  the  other  genera,  while  the  optic  lobes  of 
this  genus  with  well  developed  eyes  were  no  larger  than  in  a  Typh- 
lichthys  of  the  same  size.) 

Skeleton  not  studied.  Giinther  gives  the  vertebrae  as  thirteen  ab- 
dominal and  nineteen  or  twenty  caudal.  The  bones  of  the  head  are 
thin  and  mostly  flattened  as  in  the  Cyprinodontes.  Occiput  slightly 
convex. 

Body  compressed  posteriorly.  Head  and  anterior  portion  of  body 
depressed,  giving  the  form  of  a  broad,  flat  head,  with  a  compressed 
tail. 

Brancliiosteyal  rays  six  in  number  and  but  slightly  covered  by  oper- 
cular  bones ;  opercular  opening  large. 

Fins.  Dorsal  and  anal  nearly  opposite  and  posterior  to  centre  of 
body.  All  the  fins  except  the  ventrals  well  developed,  with  central 
rays  longest  and  first  rays  simple.  Pectorals  close  to  the  head,  about 
in  the  middle  of  the  sides.  (Ventrals  present  in  Amblyopsis,  absent 
in  Typhlichthys  and  Chologaster.) 

Mouth  opening  upwards,  with  lower  jaw  slightly  projecting.  Mar- 
gin of  the  upper  jaw  formed  by  the  intermaxillaries.  Maxillaries 
placed  behind  the  intermaxillaries,  with  lower  third  broad  and  below 
the  intermaxillaries.  Several  rows  of  fine  teeth  on  the  intermaxil- 
laries and  lower  jaw.  (Teeth  on  palatines  in  Amblyopsis  and  Typh- 
lichthys, none  on  these  bones  in  adults  of  Chologaster.) 

Scales.  None  on  the  head.  Body  closely  covered  with  small,  par- 
tially imbedded  cycloid  scales,  irregularly  arranged. 

Lateral  line  absent. 

*  From  the  Annual  Report  of  the  Peabocly  Academy  of  Science  for  1871. 

(55) 


56  SYNOPSIS    OF   THE    FAMILY   HETEROPYGII. 

Nostrils  double.  Anterior  tubular  and  standing  out  from  the  end  of 
the  snout. 

Stomach  well  defined,  coecal. 

Pytoric  appendages  present. 

Intestine  with  two  turns. 

Anus  situated  under  the  throat  and  forward  of  the  pectorals. 

Ovary  single.  (Placed  by  the  side  of  the  stomach  in  Amblyopsis 
and  Typhlichthys  and  behind  it  in  Chologaster.) 

Viviparous.     (Amblyopsis.) 

Testes  paired.     (Amblyopsis.) 

Air  bladder  with  pneumatic  duct.     (Amblyopsis.) 

Liver  with  the  left  lobe  very  large  and  partially  enclosing  the 
stomach. 

Amblyopsis  DEK.AY,  Fishes  of  New  York,  p.  187,  1842. 

Eyes  rudimentary  and  imbedded  under  the  skin. 

Head  with  numerous  transverse  and  longitudinal  rows  of  sensitive 
papilla3  provided  with  nerve  branches,  many  of  the  nerve  branches 
terminating  as  free  filaments  outside  the  papilla3.  Small  granula- 
tions on  tlie  spaces  between  the  papillary  ridges.  Canals  under  the 
skin. 

Teeth  minute,  curved,  and  arranged  in  rows  on  the  intermaxillary, 
inferior  maxillary  and  palatine  bones. 

Body  with  a  prominent  papilla  just  over  the  opercular  opening,  at 
the  base  of  a  small  papillary  ridge,  similar  to  those  on  the  head. 
Papillary  ridges  on  sides  of  body  of  same  character  as  those  on  the 
head,  and  arranged  at  nearly  equal  distances  from  opercular  opening 
to  base  of  caudal  fin. 

Pyloric  appendages,  one  on  each  side. 

Ovary  situated  on  the  right  side  of  the  stomach. 

Fins.  Ventrals  small  and  placed  near  the  anal  fin.  Dorsal,  9. 
Anal,  9.  Pectoral,  11.  Ventral,  4.  Caudal,  24. 

Amblyopsis  spelseus  DEIYAY.    LAEGE  BLINDFISH. 

CRAIGE,  Procd.  Acad.  Nat.  Sci.  Philad.,  i,  p.  175, 1842.  DEKAY,  Fishes  N.  Y.,  p. 
187, 1842.  WYMAN,  Amer.  Jour.  Sci.,  xlv,  p.  94, 1843;  Ann.  Mag.  Nat.  Hist.,  xii,  p. 
298.,  1843.  THOMPSON,  Ann.  Mag.  Nat.  Hist.,  xiii,  p.  Ill,  1844.  TELLKAMPF,  M til- 
ler's Arch.  f.  Anat.,  p.  392,1844;  N.  Y.  Jour.  Medicine,  v,  p.  84,  with  plate,  giving 
three  figs,  of  the  fish ;  position  of  internal  organs ;  brain ;  stomach ;  air  bladder;  scale 
(profile  view  gives  the  fish  without  ventral  fins,  but  ventral  view  shows  them),  1845. 
STOKER,  Synopsis  N.  A.  Fish,  p.  435,  1846.  OWEN,  Lect.  Comp.  Anat.  Fishes,  pp. 
175,  202  (fig.  of  brain),  1840.  WYMAN,  Procd.  Boston  Soc.  Nat.  Hist.,  iii,  p.  349, 1850 
"DALTON,  N.  Y.  Medical  Times,  ii,  p.  354, 18—."  AGASSIZ,  Amer.  Jour.  Sci.  xi.  p.128, 
1851.  WYMAN,  Procd.  Boston  Soc.  Nat.  Hist.,  iv,  p.  395  (1853),  1854;  v,  p.  18,  1854; 
Amer.  Jour.  Sci.,  xvii,  p.  259, 1854  (with  figs,  of  brain,  eye,  and  otolite).  GIRAKD, 
Proc.  Nat.  Sci.  Philad.,  p.  63,  1859.  POEY,  Mem.  de  Cuba,  ii,  p.  101,  Pis.  9,  11 
(outlines  of  fish  and  of  brain),  1858.  WOOD,  111.  Nat.  Hist.,  iii,  p.  314,  figure. 
1862.  TENNEY,  Nat.  Hist.,  p.  344,  figure.  1865.  GttNTHER,  Cat.  Fish  Brit.  Museum, 
vii,  p.  2, 1868.  COPE,  Ann.  Mag.  Nat.  Hist.,  viii.  p.  368, 1871.  PUTNAM,  Amer.  Nat., 


SYNOPSIS    OF   THE    FAMILY   HETEROPYGII.  57 

vi,  p.  6  et  seq.,  \vith  figs..,  Jan.,  1872.  WYMAN,  Mss.  notes  and  drawings  in  Put- 
nam, Amer.  Nat.,  vi,  p.  16  et  seq.,  1872.  PUTNAM,  Amer.  Nat.,  vi.  p.  116.  Feb., 
1872  (additional  note  on  the  young). 

PLATE  1  (American  Naturalist,  Vol.  vi,  Jan.,  1872).  FIG.  1.  Brain,  nerves  and 
organ  of  hearing  of  Amblyopsis  spelceus;  enlarged;  a,  olfactory  lobes  and  nerves; 
6,  cerebral  lobes;  c,  optic  lobes;  d,  cerebellum;  e,  organ  of  hearing,  showing  the 
semicircular  canals,  with  the  otolite  represented  in  place  by  the  dotted  lines ; /, 
medulla  oblongata ;  g,  optic  nerves  and  eye  specks.  FIG.  2.  Otolite,  enlarged. 
FIG.  3.  Eye,  magnified  (natural  size  one-sixteenth  of  an  inch  in  length);  a,  optic 
nerve;  6,  sclerotic  membrane;  c,  layer  of  colorless  cells;  d,  layer  of  pigment  cells 
(iris?);  e,  lens.  FIG.  4.  Lens,  enlarged  and  showing  the  cells.  FIG.  5.  Eye,  en- 
larged, showing  the  muscular  bands,  a,  a,  a,  a;  b,  the  lens  pressed  out  of  place;  c, 
the  optic  nerve.  FIG.  6.  Top  of  head,  showing  canals  under  the  skin,  natural 
size.  The  two  black  dots  and  lines  indicate  the  eyes  and  optic  nerves  in  position. 
FIG.  7.  Top  of  head,  showing  the  arrangement  of  the  ridges  of  papillae,  nat.  size. 
FIG.  8.  One  of  the  ridges  of  papilla?  from  the  head,  magnified.  FIG.  9.  Three  of 
the  papillae  from  the  ridge,  still  more  magnified,  showing  the  cup-shaped  summit 
and  projecting  filament.  Fig.  10.  A  portion  of  the  ridge  magnified,  and  treated 
with  acid,  to  show  the  arrangement  of  the  nervous  plexus  supplying  the  papillae 
with  nervous  filaments  from  a  branch  (a)  of  the  fifth  pair.  FIG.  11 .  Epithelial  cells 
from  the  head.  FIG.  12.  Epithelial  cells  from  the  body. 

PLATE  2.  FIG.  1.  Natural  size;  la,  stomach  and  pyloric  appendages,  twice  nat. 
size;  16,  scale,  magnified  (nat.  size  represented  by  the  small  outline  on  the  left 
over  the  figure);  Ic,  abdominal  cavity,  showing  position  of  stomach  and  single 
ovary,  nat.  size. 

Head  more  than  half  as  wide  as  it  is  long.  Length  of  head,  from 
tip  of  jaw  to  end  of  operculum,  contained  nearly  twice  in  length  of 
body  from  operculum  to  base  of  caudal  fin. 

Dorsal  and  anal  fins  of  equal  size,  rounded,  anal  commences  under 
third  ray  of  dorsal. 

Pectorals  pointed,  reaching  to  commencement  of  dorsal. 

Ventrals  pointed,  nearly  reaching  to  commencement  of  anal. 

Caudal  broad,  long  and  pointed,  membrane,  enclosing  simple  rays 
above  and  below,  continuing  slightly  on  the  tail. 

Scales  small,  longer  tljaii  broad,  with  quadrangular  centre  and  from 
8  to  12  concentric  lines,  which  are  broken  and  reduced  in  number  an- 
teriorly and  crossed  by  numerous  radiating  furrows  posteriorly.* 

Colorless,  or  nearly  so,  with  transparent  fins. 

Measurements.  Largest  specimen,  4-5  inches  total  length.  Smallest 
specimen,  1-9  total  length. 

Geographical  distribution.  Subterranean  streams  in  Kentucky  and 
Indiana. 

Specimens  examined :  — 

PROF.  WYMAN'S  COLLECTION. 

7  specimens.    Half  grown  and  adults.    Mammoth  Cave. 
MUSEUM  OP  COMPARATIVE  ZOOLOGY. 

7  specimens.    No.  778.    Half  grown  and  tf  $  adults.    Mammoth  Cave. 
1  specimen.  No. — .    Two-thirds  grown.  Cave  near  Lost  River,  Orange  Co.,  Ind. 

*T,The  scales  described  were  in  every  instance  taken  from  the  2d  or  3d  row  un- 
der the  dorsal  fin. 


58  SYNOPSIS    OF    THE    FAMILY    HETEROPYGII. 

BOSTON  SOCIETY  or  NATURAL  HISTORY. 
2  specimens.    No.  840.    Half  grown.    Mammoth  Cave. 

PEABODY  ACADEMY  or  SCIENCE. 

1  specimen.    No.  520.    Adult  9 .    Mammoth  Cave.    Presented  to  Essex  Insti- 
tute in  1851  by  N.  Silsbee. 

Other  specimens.  Dr.  Glintlier  mentions  six  specimens  and  a  skele- 
ton in  the  British  Museum.  Mr.  Thompson,  an  adult  and  newly  born 
young  in  the  collection  of  the  Natural  History  Society  of  Belfast. 
Dr.  Steindachncr  has  recently  sent  an  adult  and  eight  young  to 
the  Vienna  Museum.  The  first  specimen  of  which  we  have  any 
record  was  presented  to  the  Academy  of  Natural  Sciences  of  Phila- 
delphia ;  the  second  is  the  one  described  by  DeKay  and  then  in  the 
Lyceum  of  Natural  History  of  New  York.  Prof.  Cope  obtained  three 
specimens  from  the  waters  of  Wyandotte  Cave  in  Indiana.  Dr.  Tell- 
kampf  had  several  specimens  from  the  Mammoth  Cave,  and  it  is  prob- 
able that  specimens  exist  in  nearly  all  the  principal  museums  and  in 
many  private  collections,  as  about  all  that  have  been  caught  in  the 
Mammoth  Cave  for  years  have  been  sold  by  the  guides  to  visitors. 

Habits.  But  little  is  known  of  the  habits  of  the  large  bliudfish. 
Dr.  Tellkampf  states  that  they  are  solitary ;  on  the  slightest  motion 
of  the  water  they  dart  off  a  short  distance,  and  that  they  are  mostly 
found  near  stones  or  rocks  on  the  bottom,  and  seldom  come  to  the 
surface  of  the  water.  Prof.  Cope  states  that  if  they  are  not  alarmed 
they  come  to  the  surface  to  feed,  swim  in  full  sight,  and  can  then  be 
easily  captured  if  perfect  silence  is  preserved.  He  also  thinks  that 
they  are  principally  surface  feeders. 

In  the  stomachs  of  several  that  I  have  opened  the  only  remains 
found  were  those  of  Crayfish.  In  one  specimen,  opened  by  Dr.  Wy- 
man,  a  small  fish  with  well  developed  eyes  was  found  in  the  stomach. 
(See  Amer.  Nat.,  vi,  p.  13,  PI.  1,  fig.  13.) 

The  eggs  are  well  developed  in  September,  and  the  young  are  born 
about  the  middle  to  last  of  October.  The  young  when  born  are  half 
an  inch  or  less  in  length,  and  are  icithout  external  eyes.  (See  Amer. 
Nat.,  Feb.,  1872.  The  young  there  mentioned  may  possibly  be  those 
of  Typhlichthys.) 

Typhlichthys  GIRARD,  Procd.  Acad.  Nat.  Sci.  Philad.,  p.  C3,  1859. 

Eyes  rudimentary  and  imbedded  under  the  skin. 

Head.  The  same  arrangement  of  rows  of  sensitive  papillae  as  in 
Amblyopsis,  and  the  spaces  between  the  papillae  with  granulations  as 
in  that  genus.  (The  subcutaneous  canals  probably  exist,  but  have 
not  yet  been  made  out.) 

Teeth,  as  in  Amblyopsis,  on  the  maxillaries  and  palatines. 

Body  with  papilla  over  opercular  opening,  and  with  the  papillary 
ridges  on  the  sides  as  in  Amblyopsis. 

Pyloric  appendages    one   on  each  side   as  in  Amblyopsis,   but  of 


SYNOPSIS    OF   THE    FAMILY   HETEROPYGII.  59 

slightly  different  proportion  and  shape.  (Stomach  not  so  pointed 
behind  as  in  Amblyopsis.) 

Ovary  situated  on  right  side  of  stomach,  as  in  Amblyopsis.  (Eggs 
fewer  in  number  and  proportionately  larger  than  in  Amblyopsis.) 

Fins.  Ventrals  absent.  Dorsal,  7  or  8;  Anal,  7  or  8  ;  Pectoral,  12; 
Caudal  24.  (This  formula  is  given  after  counting  several  specimens. 
Girard  gives,  D.  7;  A.  8;  P.  11;  0.23.) 

It  will  be  noticed  that  the  only  characters  separating  this  genus 
from  Amblyopsis  are  the  absence  of  ventral  fins,  the  shape  of  the 
stomach  and  pyloric  appendages,  and  larger  eggs  in  less  number. 

Typhlichthys  subterraneus  GIKAKD.    SMALL  BLINDFISH. 

GIRARD,  Procd.  Acad.  Nat.  Sci.  Philad.,  p.  63, 1859.  GUNTHER,  Cat.  Fish  Brit. 
Museum,  vii,  p.  2,  1868  (as  a  syn.  of  Amblyopsis).  PUTXAM,  Amer.  Nat.,  vi,  p.  20 
et  seq.,  with  figs.,  Jan..,  1872. 

PLATE  2  (Amer.  Nat.,  Vol.  vi.,  Jan.,  1872).  FIG.  3,  slightly  more  than  natural  size; 
3a,  stomach  and  pyloric  appendages,  twice  nat.  size ;  36,  scale,  magnified  (nat. 
size  represented  by  small  outline  over  the  figure). 

Proportions  and  general  appearance,  want  of  color,  arrangement  of 
papillary  ridges,  position  and  shape  of  fins  as  in  Amblyopsis  spela3us, 
with  the  exception  that,  owing  to  the  jaws  being  more  obtusely  round- 
ed, the  head  is  slightly  blunter  and  broader  forward. 

Membrane  of  caudal  quite  prominent  and  extending  forwards  to  pos- 
terior base  of  dorsal  and  anal  fins. 

Scales  broader  than  long.  Large  quadrangular  centre  with  from  6 
to  8  concentric  lines  reduced  in  number  and  broken  up  on  anterior 
margin.  Posterior  portion  with  numerous  radiating  furrows. 

Measurements.  Largest  specimen,  1-85  inches  in  total  length. 
Smallest  specimen,  1*45  inches  in  total  length. 

Geographical  distribution.  Subterranean  streams  in  Kentucky,  Ten- 
nessee and  Alabama. 

Specimens  examined :  — 

MUSEUM  OF  COMPARATIVE  ZOOLOGY. 

7  specimens.    No.  780.    c?   $.   Adults.     Mammoth  Cave.    Collected  and  pre- 
sented by  Alpheus  Hyatt,  Sept.,  1859. 

1  specimen.  No.  781.    Moulton,  Alabama.    Presented  by  Thomas  Peters. 
1  specimen.  No.  782.    Lebanon,  Tennessee.    Presented  by  J.  M.  Safford. 

Other  specimens.  Dr.  Girard  described  the  species  from  a  specimen 
in  the  Smithsonian  Institution,  taken  from  a  well  near  Bowling  Green, 
Ky.  Dr.  Gunther  mentions  a  specimen,  in  the  British  Museum,  from 
the  Mammoth  Cave. 

Habits.  Nothing  is  known  concerning  the  habits  of  this  fish.  It  is 
evidently  much  rarer  at  the  Mammoth  Cave  than  the  large  species,  to 
judge  from  the  small  number  in  collections.  The  fact  that  Mr.  Hyatt 
obtained  seven  specimens  when  he  was  at  the  cave  in  September  and 
did  not  get  any  of  the  other  species,  may  indicate  some  peculiar  loca- 


60  SYNOPSIS    OF    THE    FAMILY    HETEROPYGII. 

tion  in  the  waters  of  the  cave  where  it  is  more  abundant  than  in  other 
places.  The  eggs  were  fully  developed  in  these  specimens,  but  no 
embryos  could  be  detected.  The  fish  is  probably  viviparous,  and 
very  likely  gives  birth  to  its  young  in  October. 

Chologaster  AGASSIZ,  Amer.  Jour.  Sci.,  xvi,  p.  135,  1853. 

Eyes  in  normal  position  and  well  developed. 

Head  with  small  granulations  on  the  surface  of  the  skin.  (No 
papillary  ridges.) 

Teeth  minute,  curved  and  arranged  in  rows  on  the  intermaxillary 
and  inferior  maxillary  bones.  None  on  the  palatines  in  the  adults. 
(Of  the  four  specimens  examined,  the  two  larger  (C.  cornutus)  are 
without  palatine  teeth,  while  the  single  specimen  of  C.  Agassizii, 
which  is  evidently  a  young  fish,  has  a  few  minute  teeth  on  the  pala- 
tine bones.  In  the  smallest  specimen  of  C.  cornutus  the  mouth  is 
abnormal,  the  intermaxillaries  being  reduced  to  a  small  central  portion 
and  there  are  consequently  no  teeth  in  the  upper  jaw,  but  the  minute 
teeth  on  the  palatines  are  present.*) 

{Body  without  opercular  papilla  and  papillary  ridges  on  the  sides.) 

Pyloric  appendages  two  on  each  side.  Stomach  rounded  and  turned 
slightly  on  the  side. 

Ovary  situated  principally  behind  the  stomach. 

Fins.  Ventrals  absent.  Dorsal,  8  or  9.  Anal,  8  or  9.  Pectoral,  12. 
Caudal,  28. 

This  genus  principally  differs  from  Amblyopsis  and  TyphlichthjTs  by 
the  presence  of  eyes,  the  absence  of  papillary  ridges  on  the  head  and 
body,  by  having  two  pyloric  appendages  on  each  side  instead  of 
one,  and  by  the  posterior  position  of  the  ovary.  It  agrees  with  Typh- 
lichthys  in  the  absence  of  the  ventrals,  and  the  young  further  agree 
by  the  presence  of  palatine  teeth. 

*I  believe  this  is  one  of  those  interesting  cases  where  one  set  of  organs,  or  one 
portion  of  the  animal  structure,  takes  the  place  of  another  which  from  accident  is 
wanting,  and  that  in  all  probability  these  palatine  teeth,  that  under  normal  con- 
ditions wrould  be  cast  off  as  the  fish  attained  maturity,  would  have  continued  to 
exist  in  this  specimen  and  answer  all  the  purposes  of  the  intermaxillary  teeth. 
But  that  in  this  accidental  continuance  of  these  palatine  teeth,  from  the  mere 
mechanical  use  forced  upon  them,  we  have  the  first  stages  of  the  development  of 
a  distinct  genus,  to  be  characterized  by  permanent  teeth  on  the  palatines,  and 
reduced  upper  jaw  bones,  as  many  of  the  developmental  school  would  argue,  I  do 
not  think  will  bear  the  test  of  facts  observed. 

A  not  uncommon  malformation  of  fishes  consists  in  the  entire  or  partial  absence 
of  the  maxillary  or  intermaxillary  bones.  I  have  specially  noticed  this  among  our 
common  fresh  water  trout  (Salmo)  and  marine  Conner  or  sea  perch  (Ctenolabrus) 
but  there  have  never  been  recorded  allied  genera  with  these  characters,  while 
the  malformed  specimens  are  hardly  numerous  enough  to  give  support  to  the  the- 
ory that  such  malformations  are  hereditary,  and  it  is  probable  that  each  case  was 
caused  by  the  non-development  of  the  parts  from  special  cause  during  embryonic 
life,  or  by  accident  to  the  individual. 


SYNOPSIS    OF   THE    FAMILY   HETEROPYGII.  61 

Chologaster  cornutus  AGASSIZ. 

AGASSIZ,  Amer.  Jour.  Sci.,  xvi,  p.  135,  1853.  GIRARD,  Procd.  Acacl.  Nat.  Sci. 
Philad.,  p.  63, 1859.  GUNTHER,  Cat.  Fish.  Brit.  Museum,  vii,  p.  2, 1868.  PUTNAM, 
Amer.  Nat.,  vi,  p.  21  et  seq.,  with  figs.  Jan.,  1872. 

PLATE  2  (Amer.  Nat.,  Vol.  vi,  Jan.,  1872).  FIG.  2.  Natural  size.  2a,  stomach 
and  pyloric  appendages,  twice  nat.  size.  26,  scale  magnified  (nat.  size  represented 
by  small  outline  over  the  left  of  the  fig).  2c,  abdominal  cavity  showing  stomach 
and  single  ovary  behind  the  stomach,  twice  nat.  size. 

Head  more  than  half  as  wide  as  it  is  long.  Length,  of  head,  from 
tip  of  under  jaw  to  end  of  operculum  contained  twice  in  length  of 
body  from  operculum,  to  caudal  fin.  Width  between  the  eyes  equal  to 
distance  from  eye  to  tip  of  under  jaw. 

Eyes  of  moderate  size,  situated  just  back  and  over  the  end  of  the 
maxillaries. 

Dorsal  and  anal  fins  of  nearly  equal  size,  slightly  rounded.  Anal 
with  slightly  longer  rays  and  commences  under  fourth  ray  of  dorsal. 

Pectoral  fins  pointed,  reaching  to  line  of  commencement  of  dorsal. 

Caudal  fin  pointed,  about  equal  in  length  to  the  head.  Membrane 
above  and  below  extending  but  slightly  on  the  tail. 

Scales  very  small  and  deeply  imbedded  in  the  skin.  Circular  with 
small  smooth  space  forward  of  the  centre.  From  15  to  20  con- 
centric rings,  cut  by  a  few  short  radiating  furrows  on  anterior,  and 
longer  and  more  numerous  ones  on  posterior  margin. 

Intestine  is  a  little  longer  than  in  an  Amblyopsis  of  the  same  size. 
The  two  pyloric  appendages  on  the  left  side  are  close  together  and 
broader  than  the  two  on  the  right  side,  which  are  wider  apart, 
longer  and  more  slender  than  the  others. 

Color.  Yellowish  brown,  much  darker  above,  lighter  on  sides,  and 
light  yellow  on  under  part  and  sides  of  head,  belly  and  under  part  of 
tail.  Three  longitudinal  very  dark  brown  lines  on  each  side:  the 
upper  commencing  near  the  middle  of  top  of  head  and  following 
along  the  back  to  base  of  caudal  fin ;  the  middle  one  commencing  at 
the  nostril  and  passing  through  the  eye  to  upper  portion  of  opercu- 
lum, thence  about  in  the  centre  of  side  to  about  the  centre  of  base  of 
caudal  fin ;  the  lower  commences  under  the  pectoral  fin  and  follows 
the  ventral  curve  of  the  body  to  the  base  of  caudal  fin.  All  three 
lines  are  darkest  and  broadest  forward,  and  terminate  as  a  series 
of  nearly  confluent  dots  on  the  tail.  Central  rays  of  the  caudal  dark 
brown,  outer  rays  uncolored.  Dorsal,  anal  and  pectorals  not  colored. 

Measurements.  The  three  specimens  are  respectively  1-5,  2,  and  2-3 
inches  in  total  length. 

Geographical  distribution.     South  Carolina. 

Specimens  examined: — 

MUSEUM  or  COMPARATIVE  ZOOLOGY. 

3  specimens.    No.  776.    Rice  Ditches  at  Waccamaw,  S.  C.    Presented  by  P.  C. 
J.  Weston,  1853.    (Orig.  of  Agassiz.) 


62  SYNOPSIS    OF    THE    FAMILY   HETEROPYGII. 

Habits.  Nothing  is  known  concerning  the  habits  of  this  species, 
the  only  specimens  observed  being  the  three  mentioned.  From  the 
fact  of  its  having  a  single  ovary  containing  a  small  number  (about  GO) 
of  large  eggs  it  is  probable  that  it  is  viviparous. 

Chologaster  Agassizii  PUTNAM. 

PUTXAM,  Amer.  Nat.,  vi,  p,  22  et  seq.,  with  figs.    Jan.,  1872. 

PLATE  1  (Amer.  Nat.,  Vol.  vi,  Jan.,  1872).  FIG.  4.  Natural  size;  4a,  stomach 
andpyloric  appendages,  twice  nat.  size;  46,  scale  magnified  (nat.  size  shown  lay 
minute  dot  over  left  of  the  figure). 

Head  more  than  half  as  wide  as  it  is  long.  Its  length  is  contained 
three  times  in  the  length  of  the  body  from  the  operculum  to  the  base 
of  caudal  fin. 

Eyes  proportionately  large  and  placed  over  ends  of  maxillaries. 

Dorsal  and  anal  fins  broken,  but  probably  of  about  equal  size.  Anal 
fin  commences  about  under  fourth  ray  of  dorsal. 

Pectoral  fins  pointed  and  reaching  about  half  way  to  the  dorsal. 

Caudal  fin  pointed,  not  quite  as  long  as  the  head. 

Scales  very  minute,  longer  than  wide,  with  4  or  5  concentric 
lines  round  a  granulated  centre.  A  few  radiating  furrows  cut  the 
concentric  lines  on  the  posterior  margin. 

Pyloric  appendages  and  stomach  about  the  same  as  in  C.  cornutus. 

Color.  Uniform  light  brown, without  markings  except  that  the  base 
of  the  caudal  fin  is  rather  darker  than  rest  of  fish.  Fins  uncolored. 

Measurements.    Total  length,  1-4  inches. 

Geographical  distribution.    Subterranean  streams  in  Tennessee. 

Specimen  examined: — 

MUSEUM  OF  COMPARATIVE  ZOOLOGY. 

1  specimen.    No.  777.    From  a  well  in  Lebanon,  Tenn.    Presented  by  J.  M. 
Safford.    Jan.,  1854.. 

This  species  principally  differs  from  C.  cornutus  by  having  a  longer 
body  and  smaller  head,  by  having  the  eyes  proportionately  larger,  and 
by  its  coloration.  Nothing  is  known  of  its  habits  except  the  fact 
of  its  subterranean  life.  The  scales  of  the  single  specimen  known 
indicate  a  young  fish,  and  it  is  probably  not  over  half  grown. 


The  four  species  given  in  this  synopsis  are  all  of  the  family 
as  yet  known,  but  that  others  will  be  discovered  and  the  range  of  the 
present  known  species  extended  is  very  probable.  The  ditches  and 
small  streams  of  the  lowlands  of  our  southern  coast  will  undoubtedly 
be  found  to  be  the  home  of  numerous  individuals,  and  perhaps  of  new 
species  and  genera,  while  the  subterranean  streams  of  the  central 
portion  of  our  country  most  likely  contain  other  species. 


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